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THE 



FERN 
GAZETTE 



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Edited by 

XALrabbe M.6ibby 
& B.S.Parris 



I"HE 
3R1T1SH 

PTER1D0L0GICAL 
SOCIETY 



Volume 13 Part 1 



^ 



1985 



THE BRITISH PTERIDOLOGICAL SOCIETY 

Officers and Committee for 1 985 

President: G. Tonge 
President Emeritus: J. W. Dyce 

Vice-Presidents: Dr R. E. Holttum, F. Jackson, R. Kaye, 

Prof. I. Manton 

Honorary General Secretary and A. R. Busby, University of Warwick, 

Editor of the Bulletin Science Education, Westwood, Coventry, CV4 7AL 

(Tel: Coventry 715690) 

Assistant Secretary (Membership): A. M. Paul, 

Department of Botany, British Museum (Natural History), 
Cromwell Road, London, SW7 5BD 

Treasurer: Dr B. A. Thomas, Botany Department, National Museum of Wales, 

Cathays Park, Cardiff, CF1 3NP 

Meetings Secretary: K. Kavanagh, 2 Bury Cottage, Offchurch Bury, 

Offchurch, Leamington Spa, Warwicks. 

Editors of the Fern Gazette: Dr M. Gibby, Dr B. S. Parris 

Material for publication should be sent to Dr M. Gibby, 
British Museum (Natural History), Cromwell Road, London, SW7 5BD 

Editor of the Pteridologist: M. H. Rickard, The Old Rectory, Leinthall Starkes, 

Ludlow, Shropshire, SY8 2HP 

Committee: M. Barker, J. M. Camus, P. J. Edwards, Dr N. Hards, R. P. H. Lamb, 

A. Pigott, P. Ripley, R. Rush, Dr A. Willmot, J. R. Woodhams 

Fern Distribution Recorder: A. J. Worland, 102 Queens Close, Harston, 

Cambs., CB2 5QN 

Spores Exchange Organiser: R. F. Cartwright, 13 Perry Mill Road, Peopleton, 

Pershore, Worcestershire 

Archivist: N. A. Hall, 15 Mostyn Road, Hazel Grove, Stockport, Cheshire 

The BRITISH PTERIDOLOGICAL SOCIETY was founded in 1 891 and today continues as a focus for 
fern enthusiasts. It provides a wide range of information about ferns through the medium of its 
publications and available literature. It also organises formal talks, informal discussions, field 
meetings, garden visits, plant exchanges and a spore exchange scheme. The Society has a wide 
membership which includes gardeners, nurserymen and botanists, both amateur and professional. 
The Society's journals the Fern Gazette, Pteridologist and Bulletin are published annually. The Fern 
Gazette publishes matter chiefly of specialist interest on international pteridology, the Pteridologist 
topics of more general appeal, and the Bulletin Society business and meetings reports. 

Membership is open to all interested in fern and fern-allies. Subscription rates (due on 1st January 
each year) are Full Personal Members £7; Personal Members not receiving the Fern Gazette £5; 
Student Members £5; Subscribing Institutions £8. Applications for membership should be sent to 
the Assistant Secretary (address above) from whom further details can be obtained. (Remittances 
made in currencies other than Sterling are £0.50 extra to cover bank conversion charges). 



Back numbers of the Gazette, Pteridologist and Bulletin are available for purchase from the 
Treasurer (address above), from whom further details can be obtained. 



FERNGAZ. 13(1) 1985 



A REAPPRAISAL OF DRYOPTERIS AFFINIS 

SUBSP. BORRERI VAR. ROBUSTA AND NEW RECORDS OF 

D. AFFINIS SUBSPECIES IN EASTERN EUROPE 

S. JESSEN 

Arktisch-alpiner Garten und Herpetologische Versuchsstation, 

Schmidt-Rottluff-Strasse 90, P.B. 29-07, DDR-9083 Karl-Marx-Stadt, 

East Germany 

ABSTRACT 

Dryopteris affinis var. robusta is confirmed as being a variety of subsp. borreri as 
Dryoptehs affinis (Lowe) Fraser-Jenkins subsp. borreri (Newman) Fraser-Jenkins 
var. robusta (Oberholzer & von Tavel ex Fraser-Jenkins) Fraser-Jenkins & Salvo 
Localities for var. borreri and var. robusta are listed from the author's 
collections in East Germany, Bulgaria, Czechoslovakia and Romania. In addition to 
subsp. borreri, subsp. stilluppensis also occurs as a rarity in Thuringiaand Bulgaria 
In the Northern Fagaras mountains of southern Romania var. splendens was found 
in addition to var. borreri and var. robusta; this is so far only published from 
Switzerland though Fraser-Jenkins (1 982) suggests a wider range and states that it 
may be expected elsewhere. The diploid subsp. affinis var. disjuncta, known from 
Switzerland, West Germany, Austria and Italy, was also found in southern 
Romania, thus agreeing with Fraser-Jenkins' statement(pers.comm. 1984)that he 
has also found it in Britain Spain, Portugal and France and expects it to occur 
anywhere throughout the range of subsp. affinis in sufficiently sheltered places. 

INTRODUCTION 
Fraser-Jenkins (1980, 1982) and in Dostal et al. (1984) has recognized in Europe 
(including Turkey and the Caucasus) the following subspecies and varieties of 
Dryopteris affinis (Lowe) Fraser-Jenkins: 

D. affinis 

subsp. affinis var. affinis 

var. azorica Fraser-Jenkins 

var. disjuncta (Fomin) Fraser-Jenkins 

var. punctata Oberholzer & von Tavel ex Fraser-Jenkins 
subsp. borreri (Newman) Fraser-Jenkins var. borreri 

var. pseudodisjuncta (Oberholzer & von Tavel) Fraser-Jenkins 

var. splendens (Ehrler) Fraser-Jenkins 
subsp. robusta Oberholzer & von Tavel ex Fraser-Jenkins 
subsp. stilluppensis (Sabranski) Fraser-Jenkins 

Recent information about various of the subspecies and varieties is also given by 
Eschelmuller (1972), Schneller (1974), Eschelmuller & Schneller (1980) and 
Reichstein (1983). 

Between the years 1975 and 1 984 populations of D. affinis in East Germany were 
studied by the author. Material of this species was also collected in Czechoslovakia, 
Romania and Bulgaria. In the populations studied subsp. borreri and the taxon 
described as subsp. robusta are the most frequently occurring subspecies of D. affinis. 

THE STATUS OF SUBSP. ROBUSTA 
Whereas subspp. affinis, stilluppensis and borreri are usually relatively easy to 
differentiate, distinction between subsp. borreri and subsp. robusta is often difficult 
(as mentioned by Fraser-Jenkins 1982) and arbitrary and there does not appear to be a 
clear dividing line between them. From the following observations it appears that the 
status of the taxon robusta as a subspecies of D. affinis distinct from subsp. borreri is 
incorrect and it should be a variety of subsp. borreri (and see also Table 1). 



FERN GAZETTE: VOLUME 13 PART 1 (1985) 



TABLE 1 



Features of 
distinction 



var borreri 



var. robusta 



leaves somewhat coriaceous, ± thick or 
somewhat thin, mid- to light green; 
yellowish when young; pinnule (pinna- 
lobes) -apices markedly squarely 
truncate, usually with entire or 
shallowly, squarely-lobed sides, the 
apices of the pinnules bearing + long 
acicular teeth often longer at the 
corners of the pinnule apices; indusia 
varying from relatively large at least 
when young, to small, thick or thin 
(but never as thick as in subsp. affinis), 
slightly brown to grey, lifting and 
shrivelling somewhat on ripening, often 
deciduous, usually with some being 
persistent. 



leaves herbaceous to coriaceous, 
mostly relatively thin, mid- to 
light green; yellowish when 
young; pinnule-apices rounded, 
acute or subacute, if truncate 
then only near the frond apex, 
mostly with markedly incised 
to lobed, occasionally undulate 
sides, the lobes being ± square, 
the apices of the pinnules bearing 
+ long acicular teeth; indusia 
relatively small and thin, mostly 
light grey, on ripening lifting 
and shrivelling markedly, mostly 
deciduous. 



habitat 



In Europe widespread throughout 
many habitats and altitudes; in 
East Germany occurs most frequently 
in Fagus and Fagus-combmed forests. 



the same as var. borreri. 



distribution across the whole area of the species, 

except for all of Macaronesia (Azores, 
Maderia and Canaries), and rare in 
Portugal. The present author has 
collected it in Bulgaria, Czechoslovakia, 
Romania and East Germany. Fraser- 
Jenkms (1982) gives the whole range of 
the subspecies which is the same as that 
of the variety, based on his collection 
(in BM) and herbarium specimens seen. 



the same as var. borreri. 



The two taxa are distinguishable only by the greater degree of lobing of the 
pinnules, the longer and more luxuriant pinnules and the larger frond of subsp. 
robusta (see Fraser-Jenkins 1980, 1982 and in Dostal et al. 1984); such features 
are of the type that may also occur in larger, better developed specimens of a single 
species as a result of greater growth. 

In spite of the relatively good separation between typical and well-developed plants 
of subsp. robusta (which are very frequently mistakenly identified as Dryopteris x 
tavelii Rothm.) and smaller or medium-sized subsp. borreri (also fairly often 
misidentified as D. x tavelii), transition forms between the taxa may be found in 
many populations, particularly those occurring parti ally in shade and parti ally in the 
open. 

Subsp. robusta and subsp. borreri appear not to occur growing together as distinctly 
separated taxa in the same habitat, but always to be connected by the intermediates 
mentioned. 

Cultivation of both taxa in approximately the same growing conditions show that 
the frond morphology of both taxa is able to approximate so nearly that it is difficult 
or almost impossibly artificial to tell one from the other. 

Subsp. robusta appears to be found dispersed throughout almost the whole range 
of the species (except Macaronesia and parts of North West Spain and Portugal) and 
in particular mirrors the range of subsp. borreri {see Dostal et al. 1984). Fraser- 



DRYOPTERIS AFFINIS IN EASTERN EUROPE 



Jenkins (pers. comm. 1 984) adds that this is even more markedly obvious on a local 
level, particularly in Spain and Portugal where subsp. borreri is less common than it 
is elsewhere. 

— Although requiring more study it seems fairly certain that the lack of pairing of 
chromosomes in the 16-celled sporangia resulting from the combination of 
different genomes present in subsp. robusta is the same as that in subsp. borreri 
(see Fraser-Jenkins 1982), whereas subsp. robusta and subsp. borreri show 
different pairing behaviour from subsp. stilluppensis (which shows approximately 
equal numbers of bivalents and univalents in 16-celled sporangia); subsp. affinis 
differs in being diploid. 

Fraser-Jenkins (1982) has himself come to the conclusion that subsp. robusta 
merely represents luxuriant plants of subsp. borreri, and is probably best treated as a 
variety; he also considers this, andthe other varieties treated for the purposeof Dost^l 
et al. (1 984) to be of little taxonomic importance, whereas the subspecies are far more 
significant. He points out (pers. comm. 1984) that intermediates occur between the 
varieties, and that the variation appears to arise from habitat differences, state of 
development of the plant and random vari ation genet ica I ly fixed by apomixis. Recently, 
Fraser-Jenkins and Salvo (1984) have made the new combination Dryopteris affinis 
subsp. borrero var. robusta (Oberholzer & von Tavel ex Fraser-Jenkins) Fraser-Jenkins 
& Salvo. 

Although the subspecies are of considerable taxonomic significance, neither the 
present author nor Fraser-Jenkins (per. comm.) agree with Holub (1984) who 
continues to treat D. pseudomas (Woll.) Holub & Pouzar as a species, while ignoring 
subsp. stilluppensis. He also combines subsp. robusta as a subspecies of D. 
pseudomas which presumably arose from following Fraser-Jenkins' original 
erroneous publication. It seems likely that Holub has not realised how wide-spread 
subsp. affinis is in Europe and that it was previously included under D. pseudomas, or 
that he is notfamiliar withthethreesubspecies, which aresufficiently closethat many 
botanists cannot identify them as yet. It is worth noting that Fraser-Jenkins (pers. 
comm.) also found subsp. stilluppensis in Czechoslovakia (Boscovice, N of Brno, CRFJ 
4989, 15.7.1976) and it is at present impossible to exclude the occurrence there of 
subsp. affinis as a rarity. Study of the herbarium mateial at Prague under the name D. 
pseudomas will doubtless reveal more than one subspecies. Fraser-Jenkins (in prep.) 
intends to discuss the status, distribution, distinction and other details of the sub- 
species throughout their range, but he states that it is a common misconception that 
subsp. borreri is what used to be called D. borreri or D. pseudomas, while subsp. affinis 
is a little known taxon from SW Europe and Macaronesia; in fact the most extreme 
"borreri" (as opposed to D. filix-mas) is the widespread subsp. affinis. 

NEW RECORDS OF D. AFFINIS 
The following populations of subsp. borreri (both varieties) and other subspecies of D. 
affinis have been found by the author and specimens are deposited in the herbarium of 
the Arktisch-alpiner Garten und Herpetologische Versuchsstation, Karl-Marx-Stadt. 

1. D. affinis subsp. borreri 
1.1 East Germany 
1.1-1 Thuringia 

— Eisenach: "Landgrafenschlucht", 1 specimen seen, 25.9.1977, SJ-158: var. robusta. 

— Winterstein: "Schwarzbachgraben", several specimens seen, 13.2.1977, SJ-167: var. 
robusta. 



FERN GAZETTE: VOLUME 13 PART 1 (1985) 



Suhl: "Sperbergrund" below the "Schmucke", 3-5 specimens seen, 4.10.1975, SJ-165: 

var. robusta. 

Stutzerbach near llmenau: "Marktal" (discovered by L. Meinunger), scattered. 

1 1 . 1 0. 1 979, SJ- 1 64: var. robusta. 

Steinach: "Leierloch", S slope of "Fellberg", 700m (discovered by L. Meinunger), 1 

specimen seen, 20.8.1978, SJ-162: var. robusta. 

Steinheid: "Boosgrund" NNE of Theuern, 600m (see Meinunger, 1965), 5 specimens 

seen, 1.10.1983, SJ-351: var. borreri. 

Steinheid: "Schletzenbachtal" SSW of Steinheid, 700m (see Meinunger, 1965), 2 

specimens seen, 1.10.1983, SJ-354: var. ? 

Lauscha: E slope of "Goritzberg", 640-650m (see Meinunger, 1965), ca. 5 specimens 
seen, 1.10.1983, SJ-347: var. robusta with transition to var. borreri. 
Lauscha: side valley of "Giftiggrund" S of "Pappenheimer Berg", 720m (see Meinunger, 
1965), 3 specimens seen, 1.10.1983, SJ-349: var. robusta. 

Lauscha: western side valley of "Giftiggrund" ESE of Lauscha station, 630-660m (see 
Meinunger, 1965), 10 specimens seen, 1.10. 1 983, SJ-350: var. borreri. 

Goldisthal: "Grosses Langebachtal", 650m (see Meinunger, 1965), 4 specimens seen, 

1.10.1983, SJ-352: var. borreri. 

Schalkau near Sonneberg: side valley of "Neundorfer Grund" S of Neundorf, shell 

limestone, 480m, 1 specimen seen (discovered by L Meinunger), 1.10.1983, SJ-353:var. 

robusta. 

Burgk/Saale: cistern of Burgk castle, 1 specimen seen, 20.3.1 981, SJ-368: var. robusta. 

Burgel near Jena: "Waldecker Schlossgrund", 305-325m, more than 25specimensseen, 
collected since 1975, SJ-166: var. robusta (see Fig. 1.). 

Stadtroda: wooded ravine S of Waltersdorf (see Marstaller, 1972), 3 specimens seen, 
20.1 1.1981, SJ-361: var. robusta. 




FIGURE 1 Dryopteris affinis subsp. borreri var robusta "Waldecker Schlossgrund' 
near Burgel, Thunngia (DDR), Foto S. Jessen, 1975 



DRYOPTERIS: AFFINIS IN EASTERN EUROPE 



— Stadtroda "Glastal" E of "Hermannsmuhle", 350m (see Marstaller, 1972), 4 specimens 
seen, 17.8. 1984, SJ-378: var. borreri. 

— Stadtroda "Heichbachgrund" NE of Karlsdorf, 310m (see Marstaller, 1 972), 2 specimens 
seen, 18 8. 1984, SJ-379: var. borreri. 

— Eisenberg "Grosser Tannigt" near Gruna (see Falkenberg, 1979), several specimens 
seen, 26 9.1981, SJ-364: var. robusta. 

— Gera "Teufelsgraben" SW of Reichardtsdorf (see Falkenberg, 1979), 7 specimens seen, 
26 9 1981, SJ-143: var robusta. 

— Gera: stream ravine SW of Schafpreskeln (see Falkenberg, 1979), 5 specimens seen, 
26 9.1981, SJ-362: var. robusta. 

— Gera "Turkengraben" E of Durrenebersdorf (see Falkenberg, 1979), 17.7.1982, SJ-153: 
SJ-144: var. robusta (cytologically investigated by Mrs H. Rasbach, 21.8.1984: 123 
bivalents). 

— Gera "Fuchsgraben" NW of Wolfsgefarth (see Falkenberg, 1979), 2 specimens seen, 
17.7.1982, SJ-151: var robusta. 

1.1.2 Region of the Mulde (district Leipzig) 

— Penig: sandy ground NE of Wernsdorf, 1 specimen seen, 23.6.1982, SJ-340: var. robusta 
or borreri. 

1.1.3 Dresden district ("Sachsische Schweiz") 

— Hohnstein: "Riesengrund", more than 10 specimens seen, collected since 28.3.1976, SJ- 
159: var. robusta. 

Hohnstein: "Kohlichtgrund", 2 specimens seen, 15.10.1977, SJ-160: var. robusta. 

— Hohnstein: Barenhohlgrund", 4 specimens seen, 6.8.1978, SJ-1 61 : var. robusta. 

Wehlen: "Diebskeller" ("Uttewalder Grund"), 4 specimens seen, 1 7.4. 1981, SJ-360: var. 
robusta. 

1.1.4 Halle district 

— Thale: N slope of Bode valley near "Tiergarten Hexentansplatz", 400m (discovered by L 
Meinunger), 4 specimens seen, 17.6.1984, SJ-369: var. robusta. 

1.2 Bulgaria 

1 .2. 1 Pirin mountains 

— SE Pirin: Brezniska valley NW of Goce Delcev, 1200m, 30.6.1976, SJ-1 69; var. borreri; 
1800m, 5.7.1978, SJ-1 68: var. robusta. 

— N Pirin: wood near Predel-Pass, 1200m, 1 specimen seen, 20.7.1981, SJ-331 var. borreri. 

1.2.2 Rhodope mountains 

— S of Plovdiv: ascent to "Prespa" (2000m) starting from Manastir, small wooded ravine, 
approx. 1525m, 1 specimen seen, 16.7.1983, SJ-346: var. robusta. 

1.2.3 Balkan mountains 

— Kalofer: Tuscha valley NNE of Kalofer, 1500m, 80-100 specimens seen, 16.7.1984, SJ- 
383: var. robusta. 

1.3 Czechoslovakia 

Blansko, N of Brno: stream ravine NW of Dolni Lotha, 20.7.1977, SJ-1 70: var. borreri. 

1.4 Romania 

1.4.1 N Fagaras mountains 

— Cirtisoara valley S of Cirta from 500- 1 200m, 1 5.8. 1 982, SJ- 1 38, SJ- 1 41 : var. borreri; c. 
500m, SJ-1 40: var robusta; approx. 600- 1200m, SJ- 1 39; var. splendens (noticeably 
different from other varieties). 

Serbota valley S of Porumbaca de Jos, 500- 1000m, 25.8.1982, SJ-1 33: var. borreri, var. 
splendens from 550- 900m; var robusta c. 550m in shady places. 

1.4.2 Bucegi mountains 

— W of Busteni: Alba valley, scattered, c. 950m, 27.8.1982, var. robusta and var. borreri. 

2 D. af finis subsp. stilluppensis 

2.1 East Germany, Thuringia (see Jessen, 1984). 



FERN GAZETTE: VOLUME 13 PART 1 (1985) 



— Steinach: "Schmidtsbruch" on the way to "Leierloch", about 600m, 1 2 specimens seen, 
24.8.1980, SJ-163. 

— Gera: "Grosser Tannigt" ESE of Gruna, 1 specimen seen, 26.9.1981, SJ-358. 

— Gera: "Fuchsgraben" NW of Wolfsgefarth, 260m, 1 specimen seen, 17.7.1982, SJ-145. 

— Triptis: wooded ground WNW of Hasla (see Marstaller, 1972), 4 specimens seen, 
17.10.1981, SJ-142: 

— Burgel: between "Letsche" and "Mittelberg", 1 specimen seen (see Marstaller, 1972), 
5.81984, SJ-375. 

2.2 Bulgaria, Pirin mountains 

— N Pirin: Banderica valley near "Hutte Vihren", 1890m, 21.7.1984, SJ-384. 
3 D. af finis subsp. affinis. 

3.1 Romania, N Fagaras mountains 

— side valley of "Trans-Fagaras-way" and wooded slopes of the valley near Hotel "Bilea 
Cascada", scattered between 900-1 250m, with subsp. borreri var. borreri and var. 
splendens, D. remota, D. expansa, D. dilatata, Polystichum braunii, P. aculeatum, 
Athyrium distenti folium. A. filix-femina etc; 16.8.1982, SJ-137: var. disjuncta 
(cytologically investigated by Mrs H. Rasbach, 21.8.1984: 82 bivalents). 

— Serbota valley S of PorumOaca de Jos, scattered between 650-1 200m with subsp. borreri, 
D. remota, D. expansa, D. dilatata, D. filix-mas, Athyrium filix-femina, Polystichum 
aculeatum, P. braunii. P. setiferum; 24 & 25.8.1982, SJ-1 34: by comparison with named 
specimens sent by Prof T. Reichstein this is most probably var. disjuncta; cytological 
investigation has not yet been possible. 

ACKNOWLEDGEMENTS 
I am indebted to Mr Christopher R. Fraser-Jenkins, Oxford, for suggestions and help 
with this article and for literature references, to Prof Dr T. Reichstein, Basel, for 
determination of specimens, provision of authentic herbarium specimens and for 
literature references, to Mr A. Eschelmuller, Sulzberg, for provision of material for 
comparison and for literature references, to Mrs H. Rasbach, Glottertal, for the 
cytological examination of two specimens and to Mr E. Schwerdtner, Karl-Marx-Stadt, 
for translation. 

REFERENCES 
DOSTAL, J., REICHSTEIN, T, FRASER-JENKINS, C.R. & KRAMER, K.U. 1984: Pteridophyta. In 

Hegi, G.: Illustrierte Flora von Mitteleuropa. 3. Aufi. Bd. 1, Tei/1: 1 42-1 48 Hamburg und 

Berlin. 
ESCHELMULLER, A. 1972: Dryopteris pseudomas (Wollaston) Holub & Pouzar — Typen und 

Fundorte im siidlichen Allgau. Ber. Naturf. Ges. Augsburg 27.45-65. 
ESCHELMULLER, A. & SCHNELLER, J.J 1980: Beitrag zur Kenntnis der Variability von 

Dryopteris affinis im Allgau. Mitt. Naturwiss. Arbeitskr. Kempten 24/1 : 1-12. 
FALKENBERG, H. 1979: Die Fame (Pteridophytina) des Mittleren Elstergebietes urn Gera. Verdff 

Mus. Gera 7: 5-57. 
FRASER-JENKINS, C.R. 1 980: Dryopteris affinis: a new treatment for a complex species in the 

European flora. Wil/denowia 10: 107-115. 
FRASER-JENKINS, C.R. 1982: Dryopteris in Spain, Portugal and Macaronesia. Bolm Soc. Brot. 

ser.2a.55. 175-336. 
FRASER-JENKINS, C.R. & SALVO, A.E. 1984: Sobre el genero Dryopteris Peninsula lbe>ica. 

Anales Jard. Bot. Madrid, 41 (1): 195. 
HOLUB, J. 1984. Some new nomenclatural combinations 1 . Folia Geobot. et Phytotax. 19:?AA- 

215. 
JESSEN, S. 1984: Beitrag zur Kenntnis der einheimischen Pteridophytenflora. Mitt. flor. Kart. 

Halle. 10: 76-92. 
MARSTALLER, C. 1 972: Zur Flora von Ostthiiringen (IV. Beitrag). Wiss. Z. Univ. Halle XX/72M, H 

J. 2-12. 
MEINUNGER, L. 1 965: Zur Flora von Sudthuringen (II. Beitrag). Wiss. Z. Univ. Halle XIV'65 M, H 6: 

500-502 
REICHSTEIN, T. 1983: Dryopteris affinis var punctata im Hullerich-Wald ob Pfaffikon (SZ). 

Farnblatter 9: 9-21. 
SCHNELLER, J.J. 1 974: Untersuchungen an einheimischen Farnen, insbesonderederD/yoprer/s 

filix-mas - Gruppe, 1. Ber. Schweiz. Bot. Ges. 84: 195-217. 



FERN GAZ. 13(1) 1985 



ISOLATING MECHANISMS 
IN FOUR HIMALAYAN DRYOPTERIS SPECIES 



D.S. LOYAL 
Botany Department, Panjab University, Chandigarh- 1 60014, India 

ABSTRACT 
Four diploid sexual species of Dryopteris (D. chrysocoma, D. caroli-hopei, D. 
nigropaleacea and D. cochleata) are sympatrically distributed in NW Himalaya. A 
survey of a few selected localities in Mussoorie reveals no evidence of the presence 
of interspecific hybrids there. Habitat preferences of the sporophyte, and meiotic 
and spore-release periodicity, suggest that hybridisation may be minimised due to 
these isolating mechanisms. The indusium is involved in spore-release in addition 
to its role as a protective organ. 

INTRODUCTION 
The evolutionary history of a fern genus is learnt by study of populations of its species, 
particularly of the diploid sexual species, which may betaken as representing the basic 
diversity within the genus. Four diploid sexual species of Dryopteris Adanson[D. 
chrysocoma (Christ) C.Chr., D. nigropaleacea (Fras.-Jenk.) Fras.-Jenk. (in press), 1 D. 
caroli-hopei, Fras.-Jenk. (in press), 2 and D. cochleata (Ham. ex Don) C.Chr.] show 
sympatric distribution patterns in NW Himalaya and are clearly recognisable, with 
sharp morphological discontinuities. Despite the fact that natural populations of the 
first three species coexist in the general area under investigation, inter-specific 
hybrids have not been observed there, in contrast to the well-known high incidence of 
Dryopteris hybrids in Europe and North America (for a comprehensive review, see 
Lovis 1977). It should be noted though that several workers have suggested that 
hybrids in Dryopteris are frequent between species related to each other, with 
genomes in common, but rare between unrelated species, though exceptions exist in 
the indiscriminately frequent hybrids of the N. American species, D. marginalis (L.) 
Gray. Rarity of hybrids may therefore be expected between the unrelated species 
studied in the present paper. Only one such hybrid, D. x wechteriana Fras.-Jenk. (in 
press) (= D. chrysocoma * D. nigropaleacea) has been discovered (Fraser-Jenkins in 
press, Gibby in press). The absence or at least great scarcity of hybrids also suggests 
the possibility that the chances of hybridisation and subsequent survival of hybrid 
sporophytes may be further lowered by the existence of some hitherto unknown 
isolating mechanisms. As far as the author is aware, this aspect of Himalayan fern 
biosystematics has received far less attention than it deserves. The present study 
deals mainly with habitat preferences of the sporophyte plants and with meiotic and 
spore release periodicity, in the species mentioned. 



This species is common throughout the W Himalaya where it has long been included with 
other species under the name D. odontoloma (Moore ex Bedd.) C.Chr. However following 
the discovery of two cytotypes in the complex in the Himalaya (Mehra & Loyal 1965), 
Fraser-Jenkins (in press) has separated the diploid sexual plant as D. nigropaleacea and 
the mostly E Himalayan triploid apomictic plant as D. juxtaposita Christ, while true D. 
odontoloma is a triploid apomict apparently confined to South India. 

This species is separated from D. marginata (Wall, ex Clarke) Christ by Fraser-Jenkins (in 
press) and occurs throughout the Himalaya except in the far western part, whereastrue D. 
marginata is confined to the East Himalaya. 



FERN GAZETTE: VOLUME 13 PART 1 (1985) 



MATERIAL AND METHODS 

Three populations each of D. nigropaleacea, D. caroli-hopei and D. chrysocoma 
occurring within 2-3 miles of each other were selected for study at Mussoorie in NW 
Himalaya. Several populations of D. cochleata were studied within 5 miles radius of 
Mussoorie, in the foothills of Dehra Dun. The time of meiosis was recorded during a 
series of visits to the populations commencing in June. Indusial features related to 
spore-release, such as its size, cellular changes during ontogeny and features of the 
adult indusium were studied from whole mount preparations as well as from sectioned 
material stained in an aqueous solution of Ruthenium red. The period of maximum 
spore release was recorded between July and December. 

OBSERVATIONS AND DISCUSSION 
Altitudinal and habitat isolation. Of the 4 species, D. nigropaleacea, D. caroli-hopei 
and D. chrysocoma grow in the same general localities but in single-stands or in 
isolated populations between 2000-2500m. D. chrysocoma is essentially an 
inhabitant of open, steep, usually rocky, E-facing slopes, in clearings, or beside roads, 
on well-drained soil. The rhizome is short and ascendant, bearing a rosette of closely 
set fronds. In contrast, D. nigropaleacea and D. caroli-hopei grow on shaded forest 
floors where water run-off collects somewhat plentifully due to a thick cover of 
decaying plant materials, the chief tree component being Quercus incana Bartr., 
Cornus macrophylla \Na\\.,Aescu/us indica Colebr., Ilex dipyrena Wall., Rhododendron 
arboreum Smith and Toona serrata (Royle) M.J. Roem. D. cochleata is one of only two 
species of the genus in India (the other being D. sparsa (Ham. ex Don) 0. Ktze) found in 
sub-montane forests from 800-1000m, and occurs throughout the Himalaya, except 
the far western part, and on mountains in the peninsula of India and in South India. It 
grows in different habitats from the other species, on soil along stream banks rich in 
clay or mixed clay and silt — typically an area affected by intense surface run-off. 

D. chrysocoma receives direct sunlight for the greater part of the day; D. 
nigropaleacea and D. caroli-hopei, on the other hand, are adapted to partial shade and 
weak sunlight, though some populations of the former can tolerate relatively open 
spaces which are also more dry, for example in areas of biotic disturbance such as 
where grazing or felling has occurred. In such populations the plants develop a 
somewhat stiff-textured lamina, and sporulation in these semi-xeric individuals may 
occur slightly earlier than in those occupying more shaded and less dry localities. 

As in D. chrysocoma, D. nigropaleacea has a short, ascendant rhizome bearing 
closely-set, medium-sized fronds. By contrast D. caroli-hopei has very large fronds up 
to 1 .5m and the plants inhabit more humid environments with a damp substratum. The 
rhizome is horizontal and somewhat creeping, with slightly distantly-placed fronds. 

It is clear that the sporophytes of all four species have specific ecological 
requirements which do not normally permit intermixing of the species in a given 
locality, though it may be expected that in certain transition habitats, such as the 
secondary environments at the edges of roads and clearings, they may become closely 
juxtaposed. 

Frond life-span. At the onset of the monsoon rainy season in June and July, D. 
chrysocoma produces a few vegetative fronds before the fertile ones. All the fronds 
produced during one growth season enter senescence and then die during the late 
Autumn; however they remain attached to the rhizome apex in a brown and semi- 
dried-up condition and the fertile ones release the spores during and after the Winter 
snowfall — a situation comparable with Matteuccia struthiopteris (L.) Tod. in New 
England (Klekowski, 1979). In contrast the fronds of D. nigropaleacea and D. carol- 
hopei, are biennial, i.e. the fronds may remain green for at least two years. 



ISOLATING MECHANISMS IN DRYOPTERIS 



In virtually all the fertile fronds of D. nigropaleacea examined, the soriferous 
pinnae are restricted to the upper half of the lamina; this may be connected with more 
efficient spore dispersal. D. cochleata has dimorphic fronds; the more upright fertile 
fronds have a distinctly narrower lamina and a longer stipethan the vegetative fronds. 
In all the populations examined, the vegetative fronds are produced during the 
monsoon, from July onwards and the fertile ones during October-November. 
Meiotic periodicity. Table 1 shows onset of meiosis for D. nigropaleacea in June 
during pre-monsoon showers, forD. caroli-hopei and D. chrysocoma in July during the 
monsoon, and for D. cochleata in October with low temperature, low humidity and no 
rainfall. 

Indusial features. Study of the following indusial features provides information 
relevant to the spore-release periodicity: (a) Growth rate in relation to receptacular and 
sporangial development, (b) size and cellular configurations at maturity and (c) the 
timing and extent of pectin deposition on the cell walls. 
TABLE 1. Comparison of the four Dryopteris species. 



Species 


Frond 


Onset of 
meiosis 


Maximum spore- 
release 


Indusium 


Sporangium 


Spores 


Life-span 


Size 


D. nigropaleacea 


Monomorphic, 
biennial 

Monomorphic, 
biennial 

Monomorphic, 
annual 

Dimorphic, 
annual 


June 
July 
July 
October 


Early August 

Late August 

December- 
January 

Early December 


Short-lived 
Short-lived 
Persistent 
Persistent 


874 x 600pm 

969 x 720pm 

1 150 x 890pm 

1000 x 880um 


87 x 1 10pm 

97 x 1 28pm 

160 x 200pm 

155 x 180pm 


20.5 x 16.5pm 
20 x 13.5pm 
40 x 30pm 
39 x 29pm 


D. caroli-hopei 
D. chrysocoma 
D. cochleata 



In D. chrysocoma, the prominent thick sorus remains fully closed, up to and long 
after spore formation (Figs. 1, 2). As Fig. 2 shows, D. chrysocoma has the largest 
indusium of the species investigated, with the incurved margin more-or-less reaching 
the receptacular base below the sporangia. Except for the marginal cell layers, which 
continue to divide by both anti- and periclinal divisions, all the lower-lying cell layers 
enter an expansion phase during the growth of the indusium, becoming markedly 
rectangular. A few basal, first-formed cell layers develop undulate walls which occur 
only in the oldest part of the indusium. In the other three species, virtually all the cell 
layers develop characteristic undulate walls, and once cell expansion and undulation 
have set in, further cell formation ceases. In D. chrysocoma, the indusium develops 
precociously, i.e. much earlier and faster than the subtending sporangia and its growth 
continues by cell-division and expansion for an appreciably longer period of time than 
that in the other species. An apparent advantage that accrues from this growth pattern 
is that the empty space between the developing sporangia and the indusial cover 
always contains humid, saturated air or even becomes filled with minute water 
droplets, thus preventing desiccation. Deposition of pectin on the indusial cell walls is 
meagre or, in most of the cases examined, does not even occur at all. In D. cochleata 
(Figs. 3, 4) the adult indusial form and its growth pattern is similar to that in D. 
chrysocoma, except that pectin deposition is far greater, the end result bearing a 
striking resemblance to collenchyma thickenings; the adult indusium of D. cochleata, 
though curved down at the sides, also has markedly less inflected margins which lift 
very slightly to expose the lowermost edge of the sporangial mass, on ripening. 

The thin, short-lived indusia of D. nigropaleacea (Figs. 5, 6) and D. caroli-hopei 
(Figs. 7, 8) are flap-like (more-or-less flat), or helmet-shaped (curved slightly over the 
top of the sorus), respectively. Differentiation of their constituent cells by means of 
vacuolation and expansion is completed in a relatively shorter period of time than in 
the two former species, with indusial cell-division in a sorus ceasing well before the 
development of all the sporangia in that sorus. Thus, in both these species, where the 



10 



FERN GAZETTE: VOLUME 13 PART 1 (1985) 





sis *" tdfl 


j01 


1 


2 




:/f ,,. 




FIG. 1. 0. chrysocoma, part of fertile frond FIG. 5. 0. nigropaleacea, part of fertile frond 
FIG. 2. 0. chrysocoma. indusium. FIG. 6. nigropaleacea, indusium. 

FIG. 3. cochleata. part of fertile frond. FIG. 7. caroli-hopei, part of fertile frond. 

FIG. 4. cochleata, indusia. FIG. 8. caroli-hopei, indusium. 

(Figs. 1, 3, 5, 7 Scale represents 1cm; Figs. 2, 4, 6, 8 Scale represents 0.5mm.) 



ISOLATING MECHANISMS IN DRYOPTERIS 11 



indusial margins do not curve down around the sporangia, the stage is set for spore 
release as soon as the only partially protected sporangia reach maturity during the wet 
mid to late monsoon weather of August to September. In the populations of D. 
nigropaleacea examined, maximum spore release by sporangial dehiscence occurs by 
early August. 

Spore release. Although the period during which meiosis occurs in D. chrysocoma and 
D. caroli-hopei is practically the same, spore-release in the former is postponed until 
the Winter which is a cold, dry period in the West Himalaya, or until early Spring — a 
situation akin to that in the somewhat unusual overwintering fern fronds in some 
temperate species of the genus (Farrar 1 976). Delayed spore release is associated with 
frond senescence during the period of low temperature. Fronds of D. chrysocoma 
which have not experienced senescence under low temperature, for instance 
collections made by the author in August-September in the 1950s, failed to release 
their spores despite drying as herbarium specimens. In this species both senescent 
state and the persistent indusium seem to play a role in spore-release. Apart from wind 
dispersal the snow, or water from streams passing over the rocks it often grows on, 
may carry spores or sporangia in this species. 

The possible adaptive significance of spore-release during the dry, cold season in 
D. chrysocoma and D. cochleata can be considered. The spores of D. chrysocoma (c. 40 
x 30um) and D. cochleata (c. 39 x 29pm) are distinctly larger and have a thicker perine 
than those of D. nigropaleacea (c. 20.5 * 1 6.5um) and D. caroli-hopei (c. 20* 13.5pm). 
One possible advantage of the large spore-size is that the larger quantity of food 
reserves stored in them could allow better establishment of the gametophytes before 
the rising temperatures of early summer, or could help the spores to remain alive in a 
dormant state during the considerable period they are retained on the frond in a ripe 
state in late Autumn and early Winter. It seems likely that the prothalli and sporelings 
of D. chrysocoma may be able to escape natural competition from the flush of growth of 
prothallial vegetation belonging to species from numerous genera during the 
monsoon, though further study is required into the natural growth period of its 
gametophytes and sporelings in the field. In the case of D. cochleata, the prothalli and 
sporelings (young sporophytes) may escape the intense heat anddry winds of summer 
that occur in the submontane region, and it is perhaps no coincidence that D. cochleata 
is in general a SE Asian and more southerly, lower level species, whereas the other 
species considered are all Sino-Himalayan species (see Fraser-Jenkins in press). 

THE IMPORTANCE OF ISOLATING MECHANISMS 
Finally, if it is correct that isolating mechanisms have been the cause of rarity of F1 
hybrids (as opposed to allopolyploids) in the Himalaya, the role of the different 
environments, and the past history of the Sino-Himalayan element of the genus will 
have to be considered and studied. As regards isolating mechanisms in plants in 
general, Stebbins (1950) has stated that "hybridisation between well established 
species in a stable environment will have no significant outcome — or will be 
detrimental to the species population — hence the importance of isolating 
mechanisms". The presence of numerous spontaneous hybrid plants competing with 
the parental species is clearly detrimental, and it is clear that resulting allopolyploid 
species can often out-compete the ancestral species unless the ecological 
requirements of the ancestors are so distinct that the allopolyploid species can 
themselves find an intermediate and distinct ecological niche. In Dryopteris many 
such allopolyploids (including apomicts) occur in the Sino-Himalayan region along 
with a number of sexual diploid species, all being more-or-less isolated by their 
ecology or by other factors such as those discussed in this paper. Previous reports of 
high percentages of diploid sexual species certainly do not hold true for this genus in 



12 FERN GAZETTE: VOLUME 13 PART 1 (1985) 



this region, as only two fifths of the 48 Indo-Himalayan species known at present are 
sexual diploids (see Fraser-Jenkins in press). However it is certainly true that hybrids 
of Dryopteris in the area are uncommon; Fraser-Jenkins (in press) reports nine hybrids 
from the Indo-Himalaya almost all of which are known from only one plant. One of 
these hybrids, D. x wechteriana, is between two of the species studied in this paper, D. 
chrysocoma * D. nigropaleacea (Gibby in press), and was growing as a single plant on 
a roadside above Simla, Himachal Pradesh. Analysis by Fraser-Jenkins (pers. comm. 
1983) of the kind of hybrids of Dryopteris reported from N. America (as detailed by 
Wid^n, Britton, Wagner & Wagner 1975) and from Europe (as detailed by Fraser- 
Jenkins 1982, and Dostcil et al 1984) reveals that, with certain exceptions, hybrids 
between species that are not related by having genomes in common are usually 
extremely rare, if they occur at all, whereas "backcross" hybrids, between species 
with a genome in common are frequent (see also Fraser-Jenkins, Reichstein & Vida 
1975). It would seem likely that the scarcity of hybrids in the Himalaya may be 
attributed to a similar isolation barrier between unrelated species. It does not seem 
likely that many of the seven tetraploid sexual species (a very low number)occurring in 
the Indo-Himalaya could occur sympatrically with any proposed ancestrial sexual 
diploid species, which thus reduces the possibility of triploid "backcross" hybrids 
occurring, in contrast to the situation in N.America and Europe. Nearly half of the Indo- 
Himalayan Dryopteris species are apomictic, and hybrids involving these may be 
expected to be difficult to recognise. 

ACKNOWLEDGEMENTS 
I am grateful to Mr C.R. Fraser-Jenkins, British Museum (Natural History), London, for 
help with the nomenclature and useful discussion and comments on the systematics 
of Himalayan species of Dryopteris. For the financial grant which made the field trips 
possible, I thank the Dean, University Instruction, Panjab University, Chandigarh. 

REFERENCES 
DOSTAL, J., REICHSTEIN, T„ FRASER-JENKINS, C.R. & KRAMER, K.U. 1984. Pteridophyta. In 

G. Hegi lllustrierte Flora von Mitteleuropa, 3 ed., 1: 137-192. Berlin & Hamburg. 
FARRAR, D.R. 1976. Spore retention and release from over-wintering fern fronds. Amer. Fern J. 

66: 49-52. 
FRASER-JENKINS, C.R. 1977. Three species in the Dryopteris villarii aggregate (Pteridophyta, 

Aspidiaceae), Candollea 32: 305-319. 
FRASER-JENKINS, C.R. 1982. Dryopteris in Spain, Portugal and Macaronesia. Bo/. Soc. 

Broteriana, ser. 2a, 55: 175-335. 
FRASER-JENKINS, C.R. In press. A monograph of the genus Dryopteris (Pteridophyta: 

Dryopteridacaea) in the Indian subcontinent. 
FRASER-JENKINS, C.R., REICHSTEIN, T. & VIDA, G. 1975. Dryopteris tyrrhena nom. nov. —a 

misunderstood western Mediterranean species. Fern Gaz. 11: 177-198. 
GIBBY, M. In press. Cytological observations on Indian subcontinent and Chinese Dryopteris and 

Polystichum. 
KLEKOWSKI, E.J., Jr. 1979. The genetics and reproductive biology of ferns. In A.F. Dyer (ed.), 

Experimental Biology of Ferns. Academic Press: London, New York and San Francisco: 

133-170. 
LOVIS, J.D. 1977. Evolutionary patterns and processes in ferns, in R.D. Preston and H.W. 

Woolhouse (eds), Adv. Bot. Res. 4: 229-415. 
MANTON, I. 1969. Evolutionary mechanisms in tropical ferns. Biol. J. Linn. Soc. 1: 219-222. 
MEHRA, P.J. & LOYAL, D.S. 1965. Cytological investigations in the Himalayan Dryopteris 

Adanson. Caryologia 18: 461-498. 
PAGE, C.N. 1979. The diversity of ferns. An ecological perspective. In A.F. Dyer (ed.), 

Experimental Biology of Ferns: 9-56. 
STEBBINS, G.L 1950. Variation and evolution in plants. Columbia University Press: New York. 
WIDEN, C.-J., BRITTON, DM., WAGNER, W.H. & WAGNER, F.S. 1 975. Chemotaxonomic studies 

on hybrids of Dryopteris in eastern North America. Canad. J. Bot. 53: 1554-1567. 



FERN GAZ. 13(1) 1985 13 



ANOMALOUS FRONDS AND VENATION IN 

POLYBOTRYA CERVINA 



TREVOR G. WALKER 
Department of Plant Biology, The University, Newcastle upon Tyne, NE1 7RU, 

England 

ABSTRACT 

Aberrant fronds in Polybotrya cervina are figured and described. In them the 
venation is anastomosing and complex as compared with that of normal fronds of 
this species in which the veins are simply forked and joined at the distal ends by a 
single commissural vein. 

INTRODUCTION 
Polybotrya is a genus of Caribbean, Central and South American ferns, comprising 
some 40 or more species. Most of the species are scandent on trees, despite having 
established themselves at first on the ground and with which they do not subsequently 
lose contact. Like some other scandent ferns such as Maxonia apiifolia (Sw.) C.Chr. the 
terrestrial rhizome is relatively thin but thickens considerably in its scandent phase 
(Walker 1972). Marked dimorphism of the fronds is characteristic, the sterile ones 
being simply pinnate to quadripinnate. In the fertile fronds the lamina is greatly 
reduced or absent, the segments being covered to a greater or lesser extent by 
acrostichoid sporangia. Venation of the sterile fronds is very variable, ranging from 
being completely free as in P. caudata Kze(Fig. 1 1)to completely anastomosing as in/ 3 . 
serratifolia (F£e) Kl. (Fig. 10) but without included veinlets intheareoles. A wide range 
of frond form and venation is illustrated in Tryon and Tryon (1982). 

Polybotrya cervina (L) Kaulf. is somewhat anomalous in the genus in having a 
rhizome with very limited growth with the result that the plant is not normally 
scandent but tends to grow on the forest floor, on rocks or on fallen tree trunks, etc. It is 
also very amenable to cultivation and makes an excellent pot plant. The simply pinnate 
leathery fronds (Fig. 1 ) have simple or once-forked veins which are connected at their 
distal ends by a marginal commisure (Fig. 3). The fertile fronds are more complex in 
their general architecture (Fig. 8) being bipinnate and covered with naked sporangia. 
The rather isolated position of P. cervina has led to its recognition as a distinct genus 
O/fersia by Raddi. Occasionally anomalous fronds with aberrant venation are 
produced by this species which show similarities to frond form and venation patterns 
seen in other species of Polybotrya and these form the subject of the present paper. 

ANOMALOUS FRONDS 
Anomalous fronds have arisen from time to time on plants of P. cervina from Jamaica 
and Trinidad which are in cultivation at Newcastle. These are usually photosynthetic 
fronds showing various depths of lobing and one such is shown in Fig. 2. In this 
particular frond a few sporangia were also produced on the back of the lowest pinna 
pair. A feature of such fronds is that they are of sporadic occurrence and the next 
fronds that are produced are all normal, thus the phenomenon is not genetic in the 
sense seen in the leaf 'sports' so beloved of some fern enthusiasts. However, over the 
course of years several anomalous fronds may be produced from time to time by the 
same plant. A very interesting series of specimens in the British Museum (Nat. Hist.) 
consists of a number of fronds gathered by John Smith between 1 849 and 1 859from 
a (presumably) single plant grown at Kew, some of which are illustrated here and 
showing a wide range of variation. 



14 



FERN GAZETTE: VOLUME 13 PART 1 (1985) 







FIGS. 1-5 Polybotrya cervina. Fig. 1 normal sterile frond; Fig. 2 anomalous frond; Fig. 3 normal 
venation; Fig. 4 venation in shallowly lobed frond; Fig. 5 venation in deeply lobed frond. 



ANOMALOUS FRONDS & VENATION IN POLYBOTRYA 15 



One sheet (not illustrated) shows parts of two fronds, one with normal pinnae (A) 
and the other with lobed pinnae (B) which parallel my fronds shown in Figs. 1 and 2. 
This sheet is labelled "v.v. Hort Kew 1857. A & B from the same sarmentum" 
(sarmentum refers here to the rhizome). A complete mixture of normal-shaped and 
deeply lobed sterile pinnae together with normal fertile pinnae occur together in a 
single frond gathered in 1856 (Fig. 6) whilst an 1849 specimen consists of an 
otherwise normal fertile frond in which the ends of the lowest two pairs of pinnae are 
broadened, lobed andphotosynthetic(Fig. 7); Figs. 8 and 9 show two fertile fronds from 
the same plant gathered in 1 859 in which the secondary pinnae are almost completely 
suppressed, appearing on only a few primary pinnae as undulations. 

Such anomalies as have been found at Newcastle and Kew are not merely the 
chance effects of cultivation, similar specimens having been also found in the wild as 
attested to, for example, by a specimen (Johnston 185) collected from the top of 
Tucuche in Trinidad on 29th April, 1945 (BM). 

ANOMALOUS VENATION 
One of the most remarkable features about these anomalous fronds is the effect that 
lobing of the pinnae has upon the pattern of venation. The venation changes radically 
to produce something totally different from that which might be expected to follow on a 
fairly simple change in outline of the pinna margin. Very shallow lobing such as is seen 
in Fig. 4 leads to the normal venation (Fig. 3)being more or less maintained exceptthat 
the marginal commissure is interrupted by the lobing. In the case of deeper lobing 
however such as seen in the frond illustrated (Fig. 2) the venation becomes completely 
reorganized, giving rise to a totally different pattern from that found in normal entire 
fronds. Thus, not only is each lobe provided with its own set of veins but these 
anastomose completely and also branch quite profusely giving rise to a series of 
areoles four or five deep and lacking any included veinlets (Fig. 5). This approaches very 
closely the venation pattern seen in a species such as P. serratifolia (Fig. 10), the main 
difference being that in the latter the veins are not divided up into blocks as in 
anomalous P. cervina. It should be pointed out here that there is no lobing in P. 
serratifolia to cause such a division. 

DISCUSSION 
Extrapolation from teratological forms is always a hazardous venture, especially as to 
the direction in which the change has occurred. Nevertheless, one may suggest that P. 
cervina is a simplified derived member of Polybotrya, simplified not only in frond 
morphology but also in venation. The specimens do however indicatequiteclearlythat 
in this species at least, relatively simple changes in pinna morphology leadto changes 
in venation of a most profound nature, of the order of magnitude that in most ferns 
would place two plants having the venation shown in Figs. 3and 5 in different genera. 
It points the lesson that the value of venation as a character must be separately 
assessed for each genus or group of genera and that it does not have an equal value in 
every case. 

ACKNOWLEDGEMENTS 
My thanks are due to the Keeper and Mr A.C. Jermy of the Botany Department, British 
Museum (Natural History)for facilities and providing material on whichthe silhouettes 
of the John Smith specimens are based. Thanks are also due to Miss A. T. Pickering and 
her staff for their care of ferns in cultivation at the Department of Plant Biology, the 
University of Newcastle upon Tyne. 



16 



FERN GAZETTE: VOLUME 13 PART 1 (1985) 







FIGS. 6-9 P. cervina, anomalous fronds in Herb. J. Smith, Figs. 10-11 venation patterns in 
Polybotrya. Fig. 6 collected 1856; Fig. 7 collected 1849; Fig. 8 collected 1859, normal fertile frond; 
Fig. 9 anomalous fertile frond collected from same plant as in Fig. 8; Fig. 10 venation in P. 
serratifofia; Fig. 11 venation in P. caudata. 



REFERENCES 
TRYON, R.M. & TRYON, A.F. 1982. Ferns and allied plants. Springer Verlag New York Inc. 
WALKER, T.G. 1972. The anatomy of Maxonia apiifolia: a climbing fern. Br. Fern Gaz. 10(5): 241- 
250. 



FERN GAZ. 13(1) 1985 17 



CYTOLOGY AND TAXONOMY IN WOODSIACEAE 



YI-LUN MA 
Institute of Botany, Academia Sinica, 141 Xi Zhi Men Wai Da Jie, Beijing, 

People's Republic of China 

ABSTRACT 

Chromosome counts of ten species of Chinese Woodsiaceae are reported. 
Chromosome numbers and morphological comparison support the proposal that the 
family Woodsiaceae should be divided into three genera, and the genus Woods/a 
into five sections. Base numbers of x = 33 in Protowoodsia and x = 41 , 39 and 38 in 
Woodsia may indicate relationship with Sticherus (x = 34) and Dicranopter is (x = 43) 
in Gleicheniaceae. Woodsiaceae may be derived from the common ancestor of 
Sticherus and Dicranopteris. 

INTRODUCTION 
The family Woodsiaceae includes about 30 species, mainly distributed in the northern 
temperate zone. Thirteen species (Table 2) have been previously investigated 
cytologically; the gametic chromosome numbers were found to be 41 , 39, 38, 33 or 
their multiples. The present paper provides chromosome counts of ten species from 
China, of which nine represent new counts (Table 1). 

According to Ching (1978), Woodsiaceae includes three genera, Woodsia, 
Protowoodsia and Cheilanthopsis, the latter two being monotypic with distinct 
morphological characters. Cytological evidence for this classification, and the division 
of Woodsia into Sections, is discussed. 

MATERIAL AND METHODS 
All the experimental material was collected wild in China (Table 1). Young fertile 
fronds were fixed in 1 :3 acetic acid: alcohol for 24 hours or more. Spore mother cells 
(SMCs) were examined using the squash technique of Manton (1950). Voucher 
specimens have been deposited at the Institute of Botany, Academia Sinica, Beijing 
(PE). 

RESULTS 
The chromosome counts of ten species of Chinese Woodsiaceae are summarized in 
Table 1. Photomicrographs are shown in Figures 1-12. 

1 . Woodsia lanosa Hook. A small fern distributed in SW China, over 3000m in rock 
crevices; an octoploid, n = 164. Type from Sikkim Himalaya. 

2. W. andersonii (Bedd.) Christ. A small fern distributed in W China, over 3000m in 
rock crevices. Type from Kumaon, Sikkim Himalaya. The fronds vary morphologically 
with different habitats. Cytologically it is unique: there are 1 6 SMCs per sporangium 
and 1 64 bivalents at late diakinesis, but from metaphase 1 to the formation of young 
spores meiotic division is highly irregular in most sporangia, with univalents at 
metaphase 1 and 2, lagging chromosomes and chromosome bridges at anaphase 1 
and 2. Finally, many of the sporangia contain less than 64 spores, and these are highly 
irregular in size (30-70jjm diam.) and shape. The irregular meiosis is similar to that 
seen in hybrids. The count of n = 1 64 indicates that it is an octoploid, and it may be a 
segmental alloploid derived from two closely related parents, probably W. macrospora 
C. Chr. and W. lanosa from morphological comparison. However, this needs to be 
confirmed in further study. 

*This is part of a thesis submitted as partial fulfillment for the degree of Master of Botany at the 
Institute of Botany, Academia Sinica. 



18 



FERN GAZETTE: VOLUME 13 PART 1 (1985) 
















4«* 



4* 



«?4 






•fc^ 






' * 



•v 






***** V* 1 



V 







J# tff 



FIGURES 1-12 Photographs from aceto-carmine squash preparations of SMCs at diakinesis or 
first metaphase of meiosis in ten species and two varieties of Chinese Woodsiaceae 
(magnifications in brackets). 

FIGURE 1. Woodsia lanosa n = 164 (c. 1000). 

FIGURE 2. W. andersonii n = 164 (c. 1000). 

FIGURE 3. W. rosthorniana n = 82 (c. 1200). 

FIGURE 4. W. cycloloba n = 164 (c. 1000). 

FIGURE 5. W. ilvensis n = 39 (c. 1800). 

FIGURE 6. W. polystichoides var. polystichoides n = 41 (c. 1500). 

FIGURE 7. W. polystichoides var. sinuata n = 41 (c. 1500). 

FIGURE 8a, b* W. subcordata n = c. 80 (c. 1200). 

*Figures 8b, 9b, 11b and 12b are inked in; all the rest are original photographs. 



CYTOLOGY AND TAXONOMY IN WOODSIACEAE 



19 







1* 







* ••♦ 



; 






< 



- C 










V' 10 



.'• 



*? 



** > 






ys 



12a 









Wf*i2 



FIGURE 9a, b. Protowoodsia manchuriensis n = 66 (c. 2000). 

FIGURE 10. Woodsia e/ongata n = 82 (c. 1000). 

FIGURE 1 1a, b. Cheilanthopsis indusiosa n = 74 (c. 1800). 

FIGURE 12a, b. Cheilanthopsis indusiosa showing 74 bivalents divided into two groups at 

diakinesis (c. 1200). 



20 FERN GAZETTE: VOLUME 13 PART 1 (1985) 



3. W. rosthorniana Diels. A medium-sized fern distributed in W and N China. A 
tetraploid, n = 82. Type from Sichuan. 

4. W. cycloloba Hand-Mazz. A small fern distributed in W China over 3000m, in rock 
crevices. Octoploid, n = 164. Type from the Mekong-Salwin divide, NW Yunnan. 

The above four species are all polyploids with the basic chromosome number x = 

41. 

5. W. i/vensis R. Br. A medium-sized fern widely distributed in the N hemisphere. It is 
diploid with n = 39 in the material from Heilongjiang Province, NE China. Two different 
chromosome numbers have been reported for this species, n = 41 and n = 39 (see Table 
2). It is not certain whether some of the counts are wrong, or whether there really are 
two cytotypes within this one species. 

6. W. polystichoides Eat. var. polystichoides. A medium-sized fern distributed in E 
Asia, especially throughout China. The fronds vary much in size and shape in different 
localities. A diploid, n = 41. Type from Japan. 

7. W. polystichoides Eat. var. sinuata Hook. A medium-sized fern from Liaoning, 
Shandong, and Hepei Province. Diploid, n = 41. Type from Talianwan, Liaoning. 

8. W. subcordata Turcz. A medium-sized fern from NE Asia, namely Siberia, USSR, 
Mongolia, Korea, Japan and N China. The pinnae vary in shape. Type from N China. 
There are about 80 bivalents at metaphase 1 (Figure 8; the photograph is not clear), but 
meiosis is sometimes irregular, with a few sporangia producing aberrant spores. It 
may be an allotetraploid, probably derived from W. polystichoides and W. i/vensis. 

9. W. elongata Hook. A medium-sized fern distributed in the Himalaya and SW 
China. Type from Sikkim, Himalaya. Thechromosome number has been reported as n = 
41 , diploid, by Mehra and Singh (1 955). Material collected in Yunnan has been found 
to be tetraploid, n = 82 (Figure 10). Comparison of this material with Mehra's voucher 
has not been made. 

10. Protowoodsia manchuriensis (Hook.) Ching Widely distributed in E Asia. Type 
from NE China. Kurita (1 965) found n = 33 (diploid) in Japanese material; that from 
Liaoning is tetraploid with n = 66 (Figure 9). 

11. Cheilanthopsis indusiosa (Christ) Ching A medium-sized fern found only in 
Yunnan. Type from Dali, W Yunnan. 74 bivalents are formed at diakinesis (Figure 1 1 ), 
but these are often divided into two groups of 41 and 33 bivalents in each SMC. This is 
a common phenomenon in the SMCs (Figure 12). In a few sporangia meiosis is 
irregular, and aberrant spores can be found. It may be an allopolyploid, possibly the 
bigeneric hybrid between Protowoodsia manchuriensis and Woodsia elongata, from 
cytological and morphological analysis. Further experimental study is needed. 



DISCUSSION 
Ching (1 940) established the genus Protowoodsia based on Woodsia manchuriensis. 
Now it has been confirmed that its chromosome numbers, n = 33 and 66, are different 
from species of Woodsia (n =41, 39 and 38 and their multiples). Morphologically 
there are differences, for example the sori of Protowoodsia are near the vein endings, 
and in Woodsia are situated at mid-vein; the perispore of P. manchuriensis is finely 
reticulate, unlike Woodsia. 

Thechromosome number of Cheilanthopsis indusiosa, n = 74, is different from the 
other two genera. This seems to support Ching's system (1 978), where Protowoodsia 
and Cheilanthopsis are treated as two distinct genera. Morphologically, 
Cheilanthopsis is separated by a distinct plate-like indusium, that has a few lobes that 
vary in size and shape, though it seems to be related to W. elongata in having similar 
false marginal indusia. 



CYTOLOGY AND TAXONOMY IN WOODSIACEAE 



21 



TABLE 1. Chromosome numbers for 10 Chinese species of Woodsiaceae 




Name 


Locality 


Chromosome 
numbers 


Spore 
numbers 


Ploidy 


Figure 


Voucher 
specimens* * 


1. 'Woodsia lanosa Hook 


Cangshan 

Yunnan 


n=164 


64 


8x 


1 


83 


2. 'W.andersonii 
(Bedd.) Christ 


Cangshan 
Yunnan 


n=164 


64 


8x 


2 


82 


3. 'W.rosthorn/ana Diets 


Cangshan 
Yunnan 


n=82 


64 


4x 


3 


2103 


4. 'W.cycloloba Hand-maz. 


Muli 

Sichuan 


n=164 


64 


8x 


4 


12376 


5. W.ilvensis R.Br. 


Mishan 
Heilongjiang 


n=39 


64 


2x 


5 


103 


6. W.polystichoides 

Eat. var. polystichoides 


Fenghuangshan 
Liaoning 


n = 41 


64 


2x 


6 


205 


7. 'W.polystichoides 

Eat.var.sinuata Hook. 


Fenghuangshan 
Liaoning 


n=41 


64 


2x 


7 


207 


8. 'W.subcordata Turcz. 


Fenghuangshan 
Liaoning 


n=c.80 


64 


4x 


8 


203 


9. "W.elongata Hook. 


Jizushan 
Yunnan 


n=82 


64 


4x 


10 


1651 


10 * Protowoodsia manchuriensis 
(Hook.) Ching 


Fenghuangshan 
Liaoning 


n=66 


64 


4x 


9 


202 


11. * Cheilanthopsis 

indusiosa (Christ) Ching 


Jizushan 
Yunnan 


n=74 


64 


4x 


11. 12 


157 



" first report 

** Collected by Mr. Q. Xia & Y. L Ma and determined by Prof. R. C. Ching. 

Intrageneric sub-divisions of Woods/a have differed with different authors. 
Tagawa (1937) divided the genus into four Sections: Euwoods/a Hook., Eriosorus 
Ching, Perrinia Hook, and Physematium Hook. He put stress on the presence of 
articulation on the petiole and morphology of the indusium. No cytological data were 
available at this time. Now chromosome numbers of over 20 species have been 
confirmed (Table 2). With the data available from both cytology and morphology, it is 
suggested that Woods/a be divided into five Sections (Table 3): Eriosorus, Woods/a, 
Acrolysis, Physematium and Perrinia. Within each Section there is one basic 
chromosome number, with the exception of Woodsia and possibly Perrinia. Some key 
morphological characters correspond with these Sections, for example in Sect. 
Woodsia (mostly x = 39) the articulation on the petiole is below the attachment of the 
first pair of pinnae, while in Sect. Acrolysis (x = 41 ) the articulation is at the position of 
the attachment of the first pair of pinnae. 

There has been much debate among authors concerning the phylogeny of 
Woodsiaceae. Bower (1928, 1935) considered that Woodsia is derived from 
Cyatheaceae whose phylogenetic origin could be sought in Gleicheniaceae. Ching 
(1940) also favoured a Cyatheaceae-Woodsiaceae series. Pichi Sermolli (1959) 
suggested that Cyatheales originated in the ancestors of modern Gleicheniales, and 
that Aspidiales including Woodsioideae also originated in Gleichenioid ancestors. 
From cytological data, Mehra (1961) disagreed with Bower's view; he thought that 
Woodsiaceae (x = 41) could not directly be derived from Cyatheaceae (x = 69, 70). 
Reviewing these theories, Kurita (1965) considered that n = 33 and 39 in Woodsia 
suggested affinities with n = 34 and 39 in Gleicheniaceae. 

The basic number in the most primitive group in Woodsia, Sect. Eriosorus, is x = 
41. The basic chromosome numbers in Woodsia form a series from x = 41, with 
aneuploid reduction to x = 39 and 38. The author considers that x = 41, 39 and 38 in 
Woodsia may be related to n = 43 in Dicranopteris, and that x = 33 in Protowoodsia may 
be related to n = 34 in Sticherus. Cheilanthopsis may have originated from 
hybridization between Woodsia and Protowoodsia. Woodsiaceae may be derived from 
the common ancestor of modern Dicranopteris and Sticherus. 



22 



FERN GAZETTE: VOLUME 13 PART 1 (1985) 



TABLE 2. Chromosome numbers for all species of Woodsiaceae 



Taxon 


Chromosome 
numbers 


Authority 


Woods i a R.Br. 


x=41, 39, 38 




Sect. Eriosorus Ching 


x=41 




W.macrospora C. Chr. et Max on 


no information 




W.cinnamomea Christ 


no information 




W.lanosa Hook. 


n = 164 


Y. L. Ma (present paper) 


W.rosthorniana Diels 


n=82 


Y. L Ma (present paper) 


W.andersonii (Bedd.) Christ 


n = 164 


Y.LMa (present paper) 


W.cycloloba Hand- Ma z. 


n-164 


Y.LMa (present paper) 


Sec. Woodsia 


x=39 




W.shensiensis Ching 


no information 




W.pulchella Bertol 


n = 39 2n=78 


Meyer (1959) 


W. glabella R.Br. 


n-39 2n-78 


Britton (in Fabbri 1963) (1964) 
Love, A. (1970) Love & Love (1976) 


W.hancockii Bak. 


no information 




W.ilvensis R.Br. 


n=c41 


Manton (1950) 


n-40-41 


Britton (1953) 
Taylor & Lang (1963) 


n=41 2n=82 


Love & Love (1961) Sorsa (1961, 1962) 
Vida (1965) 


n=39 2n=78 


Love & Love (1976) 
Y.L.Ma (present paper) 


W.alp'ma Gray.ssp.a/pina 


n=82 


Love & Love (1961) Vida (1965) 


n=78 2n^156 


Love & Love (1976) 


W.alpina Gray.ssp.be/li 
(Lawson) Love & Love 


n=78 2n=156 


Love & Love (1976) 


Sect. Aero lysis (Nakai) Y.LMa 
comb. nov. 


n=41 




W.macrochleana Mett. 


n=41 2n=82 


Kurita (1965) Mitui (1966, 1968) 


W.oblonga Ching et S. H.Wu 


no information 




W.pilosa Ching 


no information 




W. intermedia Tagawa 


n-82 2n = 164 


Shimura & Matsumoto (1975) 


W.polystichoides Eat.var. 
polystichoides 


n=41 2n=82 


Kurita (1961) Mitui (1968) 
Hirabayashi (1969) 
Tatuno & Okado (1970) 
Y.L.Ma (present paper) 


W.polystichoides Eat.var. 
sinuata Hook. 


n-41 


Y. L Ma (present paper) 


W.subcordata Turcz. 


n=c.80 


Y.LMa. (present paper) 


Sect. Physematium (Klf.) Hook, 
emend. 


x=41 




W.elongata Hook. 


n=41 


Mehra & Singh (1955) 


n=82 


Y. L Ma (present paper) 


Sect. Pernnia Hook. 


x=38 




W.scopulina Eat.var. 
scop/jlina 


n-38 2n=76 


Wagner (in Fabbri 1963) Brown (1964) 
Taylor & Brockman (1966) 


W.scopulina Eat. var. appal a- 
chiana (Taylor) Morton 


n = 38 2n-T76 


Brown (1964) 


W.montevidensis (Spreng.) 
Hieron. 


no information 




W.oregana Eat. 


n-38 2n-76 


Brown (1964) 


W.cathcartiana Robins. 


n=76 2n=152 


Brown (1964) 


W.obtusa (Spreng.) Terr. 


n-76 2n = 152 
n = 78 2n=156 


Brown (1964) 

Wagner & Wagner (1966) 

Wagner, Farrar & McAlpin (1970) 



CYTOLOGY AND TAXONOMY IN WOODSIACEAE 



23 



TABLE 2 (continued) 



W.plummerae Lemmon 


n=76 2n-152 


Smith, A.R. (1974) 


W.mexicana Fee 


n=c.76 2n=c. 152 


Brown (1964) 


n-82 2n-164 


Knobloch & Correll (1962) 


Protowoodsia Ching 


x = 33 




P.manchuriensis (Hook.) 
Ching 


n-33 2n = 66 


Kurita (1965) 

Mitui (1965, 1966, 1963) 


n = 66 


Y. L. Ma (present paper) 


Cheilanthopsis Hieron. 


x=74? 




C.indusiosa (Christ) Ching 


n = 74 


Y. L Ma (present paper) 


*W. mollis (Kif.) J. Smith 


no information 




*W.fragilis (Trev. ) Moore 


no information 





* Species incertae sedis. 



TABLE 3. Key to Sections of Woodsia 

1. Needle-pointed articulate hairs; petiole with or without articulation 

2. Petiole without articulation; x = 41, Sect. Eriosorus Ching in Sinensia 3: 134. 1932. 

Lectotypus W. lanosa Hook. 
2. Petiole jointed 

3. Petiole jointed below the attachment of the first pair of pinnae; x = 39 (41) Sect. 

Woodsia — based on Subsect. Ilvensis Ching. I.e. 
3. Petiole jointed at the position of attachment of the first pair of pinnae; x = 41 , Sect. 
Acrolysis (Nakai) Y. L. Ma comb. nov. — based on Subsect. Acrolysis Nakai in Bot. 
Tokyo 39: 76. 1925. Lectotypus Woodsia po/ystichoides Eat. 
1. Cylindrical or capitate hairs; petiole not jointed 

2. Presence of reflexed false indusia in addition to true indusia; x = 41 , Sect. Physematium 

(Klf.) Hook. Sp. Fil. vol. 1. 1844. Lectotypus W. elongata Hook. 
2. No false indusia, the indusium with deeply laciniate separated lobes, hidden under the 
sori; x = 38*, Sect. Perrinia Hook. I.e. Lectotypus W. scopulina Eat. 

*The chromosome numbers n = 78 in W. obtusa (Wagner, Farrar &McAlpin 1 970) and n = 82 in W. 
mexicana (Knobloch & Correll 1962) have been reported but do not correspond with n = 76 for 
these two species recorded by many other authors (see Table 2). 



ACKNOWLEDGEMENTS 
The author expresses his sincere thanks to Professors R.C. Ching and F.H. Wang for 
their help and guidance, and to Messrs Z.R. Wang and Q. Xia for varied assistance. 



24 FERN GAZETTE: VOLUME 13 PART 1 (1985) 



REFERENCES 
BOWER, F.O. 1928. The ferns. III. Cambridge, University Press. 
BOWER! F.O. 1935. Primitive land plants also known as the Archegoniatae. Macmillan & Co., 

London. 
BRITTON, D.M. 1953. Chromosome studies in ferns. Amer. J. Bot. 40: 575-583. 
BRITTOn! DM. 1964. Chromosome numbers of ferns in Ontario. Canad. J. Bot. 42: 1349-1354. 
BROWN, D.F.M. 1964. A monographic study of the fern genus Woodsia. Eastern Michigan 

University, Michigan. 
CHING, R.C. 1 940. On natural classification of the family "Polypodiaceae". Sunyatsenia 5: 202- 

' 268. 
CHING, R.C. 1 978. The Chinese fern families and genera: systematic arrangement and historical 

origin. Acta Phytotax. Sinica, 16 (3): 1-19. 
FABBRI, F. 1963. Primo supplemento alle tavole cromosomiche delle Pteridophyta di Alberto 

Chiarugi. Caryologia 16: 237-335. 
HIRABAYASHI, H. 1 969. Chromosome numbers in several species of the Aspidiaceae. J. Japan. 

Bot. 44: 113-119. 
KNOBLOCH, I.W. & CORRELL, D.S. 1962. Ferns and fern allies of Chihuahua, Mexico. Publ. Texas 

Res. Found. Renner. 
KURITA, S. 1 961 . Chromosome numbers of some Japanese ferns. II. Bot. Mag. Tokyo 74: 395- 

401. 
KURITA, S. 1965. Chromosome numbers and systematic position of the genus Woodsia. J. Japan. 

Bot. 40: 358-362. 
LOVE, A. 1 970. Islenzk ferdafldra. Reyjavik. 
LOVE, A. & LOVE, D. 1961. Some chromosome numbers of Icelandicferns and fern-allies. Amer. 

Fern J 51: 127-128. 
LOVE, A. & LOVE, D. 1976. IOPB chromosome number reports, Llll. Taxon, 25: 486-487. 
M ANTON, I. 1950. Problems of cytology and evolution in the Pteridophyta. Cambridge, University 

Press. 
MEHRA, P.N. 1 961 . Cytological evolution of ferns with particular reference to Himalayan forms. 

Proc. 48th Indian Sci. Congr. II, Presidential address: 1-24. 
MEHRA, P.N. & SINGH, HP. 1955. Cytology of Cyatheaceae, Woodsia and Marattiaceae. Curr. 

Sci. 24: 425. 
MEYER, D.E. 1959. Die Chromosomenzahl der Woodsia glabella R. Br. Mitteleuropas. 

Willdenowia 2: 214-218. 
MITUI, K. 1965. Chromosome studies on Japanese ferns. (1). J. Japan. Bot. 40: 117-124. 
MITUI, K. 1966. Chromosome studies on Japanese ferns. (2). J. Japan. Bot. 41: 60-64. 
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285-333. 
PICHI SERMOLLI, REG. 1959. Pteridophyta in W.B. Turrill (Ed.), Vistas in Botany, 7:421-493. 
SHIMURA, Y. & MATSUMOTO, S. 1975. Study on the chromosomes of Woodsia intermedia 

Tagawa. J. Geobot. 23: 2-4. 
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FERN GAZ. 13(1) 1985 25 



ECOLOGICAL OBSERVATIONS ON THE PTERIDOPHYTE FLORA 
OF LANGTANG NATIONAL PARK, CENTRAL NEPAL 

VI DYA LAXMI GURUNG 
Botanical Survey and National Herbarium, Department of Medicinal Plants, 

Kathmandu, Nepal 

ABSTRACT 

An ecological account of the pteridophyte flora of Langtang National Park of central 
Nepal is given, based on plants collected during botanical tours made during 1 977- 
1979. 142 species of ferns and 1 1 species of fern-allies in 47 genera inhabiting 
tropical to alpine regions (1230 - 4420m altitude) are reported and related to 
habitats with ecological and altitudinal data. 

INTRODUCTION 
Langtang National Park is situated in the central Himalayan region of Nepal to the 
north of Kathmandu (see Figure 1 ). The area is particularly rich in ferns. Relatively little 
attention has hitherto been given to the species of this region, and publications on its 
flora are meagre. The present report is based on botanical tours made in this area 
during 1977-1979. 

GEOGRAPHY OF THE REGION 
The area under study is from Manigaon to Langtang-Gosainkund and Helambu (parts 
of Rasuwa and Sindhupalchok districts respectively) which lie within the Park (28° to 
28°20'N latitude and 85°15' to 86°E longitude in the inner Himalayan ranges, 1230- 
4420m). From this area, extensive collections of tropicaltoalpinefernsandfern-allies 
have been made over an area of about 480 sq. miles. 

The area of the Langtang valley is of considerable scenic beauty, consisting of 
alpine meadows, lakes, glacial rivulets and the Himalayan Peaks of Langsisa (6096m) 
and Langtang Lirung (7245m). The sacred lake of Gosainkund (4420m) is the place from 
where Wallich, in 1820, procured some of the earliest gatherings of alpine plants 
made in the Himalaya. 

The character of the Langtang National Park varies from one of cool summers in 
the valley to a tundra type of climate on the peaks of the Himalayan range. 
Temperatures throughout the region also vary widely with aspect, altitude and cloud 
cover, and are generally highest from May to July and lowest from December to 
February. Precipitation in the region varies not only with altitude, but also widely 
through the year (see Table 1). The monsoon occurs between June and September, 
varying locally with topography and altitude. The area of lower valley, which receives 
quite heavy rainfall annually, also remains enveloped in thick clouds during the 
principal rainy months of June to September, when it rains almost daily and the 
humidity is at a maximum. In the upper valley, heavy downpours are generally less 
frequent. 



TABLE 1. Precipitation summary for 4stations in Langtang National Park for the years 
1976-1980 (from data kindly supplied by the meteorological service of His 
Majesty's Government of Nepal) 

DHUNCHE TARKEGHYANG SHARMATHANG TIMURE 

Lowest minimum 6mm (Nov Dec 1975) 1mm (Nov 19781 3mm (Nov 19761 limn (Jdn 1977) 

Highest maximum 649mm (Aug 1975) 763mm (July 19781 1524mm (July 1978) 301mm (July 19781 

Lowest annual amount 1661mm (1977) 772mm(1975) 3576mm(1976) 252mm(197bl 

Highest annual amount 2433mm (1979) 2668mm(1978) 4975mm(l978) 1123mm (19781 
Maximum in 24 hrs. 100mm (8 June 1976) 96mm (2 June 19 78) 1 42mm (24 July 1979) 69mm (13 Sept 1977) 



26 



FERN GAZETTE VOLUME 13 PART 1 (1985) 



FIGURE 1. Map of Langtang National Park. 




The soil of the hills and mountains in the upper valley above 3000m is composed 
mainly of sandy loam with a large proportion of gravels. In the lower forested region 
(2000 - 3000m) there is a covering humus layer of ,2 - 10cm depth. This serves as a 
good substratum for the growth of terrestrial species. Below 1 000m there are also red 
and yellow clays 



FERNS OF LANGTANG NATIONAL PARK 27 



GENERAL VEGETATION 
On the way to Langtang National Park the following forest types have been observed. 
At the lowest elevations (up to 1000m) the area carries a tropical Low Hill Forest, 
dominated by trees of Shorea robusta. Between the area of Manigaon to Dhunche 
(1 230-2000m) most of the mountain slopes have been cleared of forest for cultivation 
and a number of encroachment areas are observed. Consequently only a few thick 
forests are seen, representing the original sub-tropical Middle Hill Forest. Pinus 
roxburghii, Schima wallichii, Castanopsis indica, C. t;ribuloides and Lyonia are the 
most striking trees of the forest. Between the area of Khangjing and Ghore Tabela 
(2000- 3000m) elevation, is temperate Upper Hill Forest. This is a damp anddark ever- 
green Oak and mixed broadleaved forest composed of Quercus semecarpifolia, Q. 
lanuginosa, Q. g/auca, Q. lamellosa, Q. incana, Pinus wallichiana and Tsuga dumosa. 
To the east of the gorge of Trisuli River are Tsuga covered hills. With increasing 
altitude, Rhododendron arboreum, R. campanulatum and Alnus nepalensis occur 
luxuriantly and the forest becomes gloomier. The Oaks here bear branched crowns 
and are profusely covered with mosses and epiphytic ferns. The forest is thick and 
moist. The shelter of the trees provides shade and helps to prevent rapid air 
movements to a considerable extent, resulting in a prolific growth of small herbs, 
shrubs and ferns on the forest floor. Higher up there is gradual replacement of Oaks by 
Rhododendron, marking the beginning of the sub-alpineRhododendron-Conifer Zone. 
In the area from Ghore Tabela to Langtang village (3000 - 4000m) on the north-facing 
mountain slopes is preserved a very rich collection of conifers in a forest composed 
mainly of Abies spectabilis, Betula utilis and Rhododendron arboreum var. cambellii. 
The Himalayan Silver Fir , Abies spectabilis, forest is locally a very striking feature, from 
Chandabari and Laurebinda. The south-facing mountain slopes are partly grass- 
covered, but also have Larix himalaica, dwarf bushes of Rhododendron lepidotum. 
Juniper us recurva. Allium wallichii. Ephedra jerardiana, Rosa macrophylla, Caragana 
sp. and scrub of Cotoniaster. Betula utilis is found in the high level Rhododendron 
forest of the southern mountain range of the upper Langtang valley. At heights above 
4000m on exposed sunny slopes there is often a mosaic of Rhododendron 
campanulatum or scrub of Rhododendron lepidotum, R. anthopogon and Caragana sp. 
with Aconitum beneath. Grass-lands are also frequent in this area or above, giving 
way eventually to rocky moraine with scattered grasses and herbs. 

EPIPHYTIC FERNS 
The composition of the epiphytic element changes markedly with altitude as well as 
the shady or exposed nature of the forest. Certain species like Shorea robusta in the 
lower hill forest and the conifers at higher elevations do not afford shelter for many 
epiphytic ferns. 

Due to the relative high temperature and low humidity the epiphytes of the low hill 
zone (up to 1000m) have developed marked features for storing water. Common 
species on Shorea robusta are Lepisorus nudus and Pyrrosia mollis, in which the 
fronds roll inwards from their margins to conserve water during dry weather. 

Epiphytic ferns in the middle hill zone (1 000 - 2000m) are mostly influenced by the 
monsoon. During this season they appear prolifically, complete their annual cycle 
and their fronds wither. The rhizomes remain dormant to renew their activity the 
following year. Due to the prevailing warm temperature, adequate rainfall and high 
humidity, the branches and trunks of trees are often thickly covered with mosses and 
liverworts which serve as a sponge for water absorption, helping protect the rhizomes 
of epiphytic ferns from drought. The common epiphytes growing mainly on Schima 
wallichii and Castanopsis indica in rather open sites are Asp/enium laciniatum. 



28 FERN GAZETTE VOLUME 13 PART 1 (1985) 



Arthromeris wallichiana, Drynaria mollis, Crypsinus hastatus, Polypodium 
lachnopus, Pyrrosia mollis and Oleandra wallichii. Oleandra wallichii possesses a 
strongly branched rhizome which gives off stiff rhizome-like roots into the humus of 
the tree trunks. In more shaded areas of trunks of Castanopsis indica, C. tribuloides or 
Lyonia sp. are Asp/enium ensiforme, Lepisorus nudus, L kashyapii, Polypodium 
amaurolepida, P. amoenum, P. argutum and P. microrhizoma. 

The Quercus-Rhododendron Zone (2000 - 3000m) also has favourable conditions 
for epiphytic growth. Some of the commonest species on the trunks of Quercus 
semecarpifolia and Q. lanuginosa in the more open parts of this area are Dava/lodes 
membranulosum, Crypsinus ebenipes, Drynaria mollis, Lepisorus clathratus and L 
kashyapii. In shaded and more moist parts of the area, mostly on the trunks of Quercus 
glauca, Q. lamellosa, Q. incana. Rhododendron arboreum, R. campanulatum and 
Alnus nepalensis, there are Araiostegia c/arkei, A. de/avayi, A. pu/chra, Leucostegia 
immersa, Arthromeris himalayensis, Crypsinus oxy/obus, C quasidivaricatus, 
Drynaria propinqua, Lepisorus lor if or mis, Polypodium argutum and Pyrrosia mollis. 
Another conspicuous fern is Vittaria f/exuosa, hanging from lower tree trunks. 

In the Rhododendron-Conifer zone the epiphytic ferns decrease above 3000m. At 
the top of ridges the trees are rather short and stunted due to direct exposure to strong 
winds, but during the monsoon they receive plenty of rain and their branches are 
almost continuously immersed in fog. As a consequence they are thickly clothed with 
liverworts and mosses, forming a substrate for prolific growth of Polypodium 
lachnopus especially on Rhododendron arboreum var. campbellii. In shady areas 
Araiostegia de/avayi and Lepisorus kashyapii are seen more frequently. 

TERRESTRIAL FERNS 
At lower elevations under 1000m the forest floor is fairly poor in humus andthe soil is 
composed of reddish -brown clay. Here growing abundantly are Adiantum caudatum, 
A. philippense, Onychium siliculosum, Pityrogramma calomelanos and Sphenomeris 
chinensis. In partly shaded places Tectaria macrodonta and Pteris dactylina become 
predominant. Pteris aspericaulis and Athyrium acrostichoides frequently colonise the 
more open forest borders and Equisetum debile occurs on gravelly soil especially at 
waysides. 

A vast number of ferns grow on the forest floor between 1 000 - 2000m. The most 
frequent are Diplazium multicaudatum, D. esculentum, Athyrium macrocarpum, A. 
pectinatum, Dryopteris atrata, D. cochleata, D. marginata, D. panda, Polysticum 
lentum, P. squarrosum, Thelypteris auriculata, T. extensa, T. ornata, T. paludosa, 
Botrychium lanuginosum, Mecodium exsertum, Woodwardia unigemmata and 
Arthromeris wallichiana. On dry and exposed places along forest borders 
Dennstaedtia appendiculata often flourishes with Polystichum prescottianum, 
Onychium lucidum, Lepisorus nudus, Microsorum membranaceum, Polypodium 
amoenum, Pyrrosia flocculosa, Vittaria f/exuosa, Selaginella monospora, S. subdiaphana, 
Mecodium exsertum, Leucostegia delavayi, L. immersa and Nephrolepis cordifolia. 

At higher elevations, between 2000 - 3000m on the forest floor a large number of 
terrestrial ferns form an exuberant growth in the shade of trees and shrubs. The more 
common ferns are Pteris aspericaulis, P. vittata, Acrophorus stipellatus, Athyrium 
macdonnelli, A. macrocarpum, Dryopteris paleacea, Thelypteris erubescens, 
Cheilanthes grisea, Coniogramme caudata and C. fraxinea. The borders of the forest 
are covered with many plants of Dennstaedtia appendiculata, Leucostegia immersa, 
Athyrium filix-femina, A. foliolosum, A. japonicum, A. setiferm, Diplazium stoliczkae, 
Dryopteris chrysocoma, D. marginata, D. odontoloma, Peranema cyatheoides, Pteris 
biaurita, P. cretica, P. excelsa, P. aspericaulis, Polystichum lentum, P. 



FERNS OF LANGTANG NATIONAL PARK 29 



nigropaleaceum, P. squarrosum, P. stimulans, Thelypteris arida, T dentata, T 
esquirolii. Woodward/a unigemmata, Asplenium ens if or me, A. indie urn, Arthromeris 
himalayensis, Lepisorus lor if or mis, Microsorum membranaceum, Lycopodium 
hamiltonii, Selaginella chrysocaulos and S. subdiaphana. A characteristic fern of the 
margins of this forest is Pteris wallichiana, which possesses a large digitately 
compound frond springing from an ascending rhizome. On moist gravelly soils or in 
damp places along waysides or water channels near the forest margins Equisetum 
diffusum grows in profusion. 

The most significant undergrowths of Quercus-Rhododendron forest are formed 
by Athyrium acrostichoides, A. macdonellii, A. macrocarpum and A. n /'gripes. This 
community can cover a vast area of the forest floor between 2700 - 3100m. 

On forest borders or in open places the Athyrium community is replaced by 
Dennestaedtia appendiculata, Pteris cretica, P. quadriaurita, Araiostegia clarkei, 
Dryopteris brunoniana, D. chrysocoma, Diplazium multicaudatum, Polystichum 
nepalensis, P. nigropaleaceum, and Thelypteris paludosa. On the higher limits of the 
Quercus-Rhododendron forest (3000 - 4000m) Dryopteris brunoniana and 
Polystichum prescottianum become prominent on the forest floor or along forest 
margins. 

THICKET-FORMING SPECIES 
At elevations of up to 2000m Dicranopteris linearis is a characteristic species on 
exposed ridges of ravines, open rocky hills and on gravelly soil along waysides. It is 
exposed to wide variations of temperature and is able to tolerate direct sun as well as 
moist cool winds. It is a good soil binder as it forms thickets, but it seems to exclude all 
other species from growing in the area it occupies. Gleichenia glauca forms extensive 
thickets between 2000 - 3000m. The young fronds bear a pair of opposite pinnae 
whilethe apex becomes dormant and is closely surrounded by scales. Inthe following 
year another pair of pinnae is formed by the activation of the terminal bud of the frond, 
after which it becomes dormant again. This process is repeated year after year until 
the fronds reach huge dimensions. Pteridium aqui/inum occurs extensively over the 
area, forming huge dark green thickets in open places between 1 230 - 3400m. The 
fronds are large and the rhizomes deep underground. It is one of the species that 
sprouts vigorously after burning. Thus these tough ferns play a useful role in reducing 
soil erosion and landslides. 

CLIMBERS 
Most climbers are in the lower zones; for example Lygodium f/exuosum and L. 
japonicum are fairly common in the open areas as well as on the forest floor between 
about 1 230 - 1 900m. They grow on h-umus rich soil and the rachides of the fronds 
twine around neighbouring shrubs or branches of trees, enabling them to climb to 
great heights and into bright lights conditions. 

RAVINE FERNS 
This category includes all the fern species which grow in ravines along water 
channels on calcareous stony soil. At lower altitudes, to about 1000m, Ceratopteris 
thalictroides occurs scattered along sandy or clayey edges of water channels. On 
moist rocks along the streams at higher elevations (2000 - 3000m), are found 
Dryopteris marginata, Polystichum aculeatum, Thelypteris auriculata, Asplenium 
ensiforme, Drynaria prop in qua, Lepisorus kashyapii, Microsorum membranaceum, 
Crypsinus malacodon and Polypodium lachnopus. Lycopodium c/avatum also grows 
near water courses with Equiesetum debi/e. 



30 FERN GAZETTE VOLUME 13 PART 1 (1985) 



LITHOPHYTES 

True rock ferns 

The species that colonise dry boulders and rock crevices at low altitudes (1000m) 
include Cheilanthes dalhousiae, C. albomarginata, C. tenuifclia, Pityrogramma 
calomelanos, Onychium siliculosum, Adiantum philippense and Sphenomeris 
chinensis. Between 1 000 - 2000m the more common species are Pteris aspericaulis, 
P. wallichiana, Araiostegia pulchra, D aval lodes membranulosum, Athyrium 
drepanopterum, Dennstaedtia appendiculata, Mecodium crispatum, Diplazium 
giganteum, Hypodematium crenatum, Asplenlum laciniatum. Arthromeris 
hima/ayensisandA. wallichiana, and associations of these species can be particularly 
abundant. Less frequent are Crypsinus oxylobus, Drynaria propinqua, Loxogramme 
invo/uta, Microsorum cuspidatum, M. membranaceum, Lycopodium hamiltonii and L 
ulicifolium. 

Growing luxuriantly on shady and moist rocks within the forest between 2000 - 
3000m altitude are Adiantum venustum, Cheilanthes anceps, Dryopteris odontoloma, 
Polystichum aculeatum, P. neolobatum, P. obliquum, P. squarrosumandP. stimulans. 
Occasionally large plants of Woodwardia unigemmata occur here, and large species of 
Asplenium including A. ens/forme, A. laciniatum and A. varians. 

Many epiphytes also grow as rock ferns on moist, shady rocks within the forest 
because of the mild, moist, humid conditions. The dominant species are Crypsinus 
oxylobus Drynaria mollis, D. propinqua, Lepisorus nudus, L. scolopendrium, 
Crypsinus hastatus, Polypodium amoenum, P. lachnopus and Vittaria f/exuosa. Less 
frequent are Se/agine/la involvens, S. pennata, S. subdiaphana, Doryopteris concolor 
and Adiantum venustum, especially on dry exposed rocks along waysides. 

Between 3000 and 4000m Woodsia e/ongata luxuriantly colonises exposed dry 
rocks. Other abundant species are Cheilanthes albomarginata, C. dalhousiae, C. 
subvillosa, Cryptogramma crispa, Polystichum atkinsonii, P. prescottianum, 
Jhelypteris e/wesii, Lepisorus kashyapii, Crypsinus malacodon, Dryopteris 
chrysocoma and Pteridium aquilinum. 

Above 4000m Dryopteris brunoniana and Polystichum prescottianum appear. 

Ferns of stone-wall crevices and gravel embankments 

The main xerophytic ferns that thrive well in these dry and exposed habitats between 
1000 - 2000m are Pteris biaurita, P. vittata, Jhelypteris torresiana and Tectaria 
macrodonta, whilst Nephrolepis cordifolia may cover enormous areas locally. This last 
species possesses thin, wiry, widely creeping, branched runners which sprout at 
invervals to give rise to new individuals. The branches of these runners bear whitish 
edible water tubers at their ends. Under extremely dry conditionsthefrondsshedtheir 
pinnae whilst the rachides persist. Less abundant species are Osmunda c/aytoniana, 
Pteris nepa/ensis, Leucostegia immersa, Hypodematium crenatum, Jhelypteris 
molliuscula, Microsorum membranaceum, Polypodium lachnopus, Lycopodium 
clavatum and Se/agine/la involvens. 

At the higher elevations (2000 - 3000m) the number of ferns diminishes in this 
habitat, but Botrychium lanuginosum, Dryopteris chrysocoma, Dennstaedtia 
appendiculata and Oleandra wallichii can be found. Above 3000m, only Crypsinus 
ebenipes and Cryoptogramma brunoniana are frequent, but above 4000m elevation 
only Cryptogramma brunoniana is present. 

PHYTOGEOGRAPHICAL COMPARISON 
The pteridophyte flora of Langtang National Park, central Nepal resembles both the 
east and west Nepal floras (see Table 2). Analysis of the distribution of the species in 
Nepal is as follows: — 



FERNS OF LANGTANG NATIONAL PARK 



31 



a. No. of species from Langtang National Park, central Nepal 153 

b. No. of species common with east Nepal 1 25 (81 .69%) 

c. No. of species common with west Nepal 60 (39.21%) 

d. No. of species found only in Langtang National Park 19(12.41%) 

The above comparison shows that the pteridophyte flora of Langtang National Park has 
affinities with the floras of both east and west Nepal, although there are more species 
in common with those of east Nepal (81 .69%), than with those of west Nepal (39.21 %). 

ACKNOWLEDGEMENTS 
I am very grateful to Dr SB. Malla, Director General, Dept. of Medicinal Plants for 
kindly giving me the opportunity for the collection and field-study of ferns in relation to 
vegetation in Nepal, and to Dr S.B. Rajbhandari, Deputy Director General for his 
valuable suggestions and encouragement. 



TABLE 2. Phytogeographical distribution of ferns in Nepal Himalaya in relation to 
Langtang National Park region. 

Key to distributions: 

1 West Nepal (the area lying on 80° to 83°E longitude). 

2 Central Nepal (the area lying on 85°15' to 86°E longitude). 

3 East Nepal (the area lying on 86°30' to 88°10'E longitude). 



Scientific name 


Distribution 


Scientific name 


Distribution 


Botrychium lanuginosum 


1 


2 




C. affinis 


1 


2 


3 


Osmunda claytoniana 


1 


2 


3 


C. fraxinea 




2 


3 


Lygodium flexuosum 


1 


2 


3 


Cryptogramma crispa 


1 


2 


3 


L. japonicum 




2 


3 


Dennstaedtia appendiculata 




2 


3 


Dicranopteris linearis 




2 


3 


Doryopteris concolor 




2 


3 


Gleichenia glauca 




2 


3 


Onychium contiguum 


1 


2 


3 


Mecodium badium 


1 


2 


3 


0. japonicum 




2 


3 


M. crispatum 


1 


2 


3 


0. lucidum 




2 




M. exsertum 




2 


3 


0. siliculosum 




2 


3 


Actiniopteris semiflabellata 




2 


3 


Pityrogramma calomelanos 




2 


3 


Adiantum caudatum 




2 


3 


Pteridium aquilinum 


1 


2 


3 


A. edgeworthii 


1 


2 




Pteris aspericaulis 




2 


3 


A. philippense 


1 


2 


3 


P. biaurita 


1 


2 


3 


A. venustum 


1 


2 




P. cretica 


1 


2 


3 


Cheilanthes anceps 




2 


3 


P. dactylina 


1 


2 


3 


C. albomarginata 


1 


2 


3 


P. excelsa 


1 


2 


3 


C. dalhousiae 


1 


2 


3 


P. geminata 


1 


2 




C. farinosa 


1 


2 


3 


P. nepalensis 




2 


3 


C. grisea 




2 




P. quadriaurita 


1 


2 


3 


C. rufa 


1 


2 


3 


P. vittata 


1 


2 


3 


C. subvillosa 




2 




P. wallichiana 


1 


2 


3 


C. tenuifolia 




2 


3 


Sphenomeris chinensis 




2 


3 


Coniogramme caudata 




2 


3 


Ceratopteris thalictroides 


1 


2 





32 



FERN GAZETTE VOLUME 13 PART 1 (1985) 



Scientific name 


Distribution 


Scientific name 


Distribution 


Araiostegia clarkei 




2 




T. erubescens 




2 


3 


A. dareiformis 




2 


3 


T. esquirolii 




2 




A. delavayi 




2 


3 


T. extensa 




2 




A. pulchra 


1 


2 


3 


T. molliusculla 




2 


3 


Davallodes membranulosum 




2 


3 


T. ornata 




2 


3 


Leucostegia immersa 


1 


2 


3 


T. paludosa 




2 


3 


Nephrolepis cordifolia 




2 


3 


T. torresiana 




2 


3 


Oleandra wallichii 


1 


2 


3 


Woodsia elongata 


1 


2 




Acrophorus stipellatus 




2 


3 


Woodwardia unigemmata 


1 


2 


3 


Athyrium acrostichoides 




2 




Asplenium ensiforme 


1 


2 


3 


A. drepanopterum 




2 


3 


A. indicum 




2 


3 


A. filix-femina 


1 


2 


3 


A. laciniatum 




2 


3 


A. foliolosum 


1 


2 




A. unilaterale 




2 


3 


A. macdonnellii 




2 


3 


A. varians 


1 


2 


3 


A. macrocarpum 




2 


3 


Arthromeris himalayensis 




2 


3 


A. nignpes 




2 


3 


A. wallichiana 


1 


2 


3 


A. pectinatum 




2 


3 


Crypsinus ebenipes 




2 


3 


A setiferum 




2 


3 


C. oxylobus 




2 


3 


A. tenellum 




2 




C. quasidivaricatus 




2 


3 


Diplazium esculentum 


1 


2 


3 


Drynaria mollis 




2 


3 


D. giganteum 




2 




D. propinqua 


1 


2 


3 


D. multicaudatum 




2 




Lepisorus clathratus 




2 


3 


D stoliczkae 


1 


2 


3 


L. excavatus 




2 




Dryoptens acutodentata 




2 


3 


L. kashyapii 




2 


3 


D apiciflora 


1 


2 




L. loriformis 


1 


2 


3 


D. atrata 


1 


2 


3 


L. nudus 


1 


2 


3 


D barbigera 


1 


2 


3 


L. scolopendrium 




2 


3 


D brunoniana 


1 


2 


3 


Loxogramme involuta 


1 


2 


3 


D chrysocoma 


1 


2 


3 


Microsorum cuspidatum 




2 


3 


D cochleata 




2 


3 


M. membranaceum 




2 


3 


D marginata 




2 


3 


Crypsinus hastatus 




2 




D. odontoloma 




2 


3 


C. malacodon 


1 


2 


3 


D. paleacea 




2 


3 


Polypodium amaurolepida 




2 




D panda 




2 


3 


P. amoenum 




2 


3 


D. sinofibnllosa 




2 


3 


P. argutum 


1 


2 


3 


Elaphoglossum petiolatum 




2 


3 


P. atkinsonii 




2 


3 


Hypodematium crenatum 




2 




P. lachnopus 


1 


2 


3 


Peranema cyatheoides 




2 


3 


P. microrhizoma 


1 


2 


3 


Polystichum aculeatum 




2 




Pyrrosia floculosa 


1 


2 


3 


P. atkinsonn 




2 


3 


P. mollis 


1 


2 


3 


P. lentum 


1 


2 


3 


Vittaria flexuosa 




2 


3 


P. neolobatum 




2 


3 


Lycopodium clavatum 




2 


3 


P nepalense 




2 


3 


L. hamiltonii 




2 




P. nigropaleaceum 




2 


3 


L ulicifolium 




2 




P obhquum 


1 


2 




Selaginella chrysocaulos 




2 


3 


P prescottianum 


1 


2 


3 


S. involvens 




2 


3 


P. squarrosum 


1 


2 


3 


S. monospora 




2 


3 


P. setiferum 




2 


3 


S. pennata 




2 


3 


P. stimulans 


1 


2 


3 


S. subdiaphana 




2 


3 


Tectana coadunata 




2 




S. vaginata 




2 


3 


T macrodonta 


1 


2 


3 


Equisetum debile 


1 


2 


3 


Thelyptens arida 


1 


2 


3 


E diffisum 




2 


3 


T aunculata 




2 


3 










T dentata 




2 


3 










T. elwesn 




2 













FERN GAZ. 13(1) 1985 33 



NISTARIKA, A NEW GENUS OF POLYPODIACEAE 
FROM SILENT VALLEY, SOUTH INDIA 



B.K. NAYAR, P.V. MADHUSOODANAN and M.J. MOLLY 
Botany Department, Calicut University, 673 635, Kerala, India 

ABSTRACT 

A new genus, Nistarika, with one species, N. bahupunctika, is described, 
characterised by having four rows of simple fronds restricted to the dorsal surface 
of the rhizome, and lamina-less fertile fronds bearing sporangia directly on the 
midrib in two lateral linear sori. The genus is related toZ.epfoc/7/'/tvs(Polypodiaceae) 
and is possibly derived from Microsorum. 

INTRODUCTION 
Silent Valley, a small, rather densely forested area of the Western Ghats of south 
India close to the Palghat Saddle on the western fringe of Nilagiri ranges, has recently 
attracted attention as one of the few undisturbed wet evergreen tropical forests left in 
the Indian subcontinent. It is a small table-land on the western slope of Western 
Ghats, criss-crossed by many fastflowing rivulets which havecutdeep intotheterrain 
making it highly dissected. The slopes are covered by dense wet evergreen forests and 
in many cases the ridge crests are covered in grassland. Undergrowth is sparse inthe 
wooded regions, except on the banks of streams and borders of grassland and other 
open areas. The climate is equatorial with a temperature range of 15-25°C, and an 
annual rainfall of c. 380cm, mostly from May to September. Even during the dry 
season (January-May) the forest floor never becomes exposed. As Silent Valley was 
botanically unexplored on account of its inaccessibility and the rugged terrain, a 
f loristic study was undertaken of the fern flora and over 1 00 taxa recorded from the c. 
2000ha studied (Nayar 1982). One of the ferns collected appears to be as yet un- 
named; it is described here together with some aspects of its morphology. It grows in 
dense shade at forest margins and is fairly widespread in the Panthanthodu and 
Kummattanthodu areas but is less abundant on the banks of Kunthippuzha River in 
the southern half of Silent Valley Reserve forest. 

TAXONOMIC DESCRIPTION 
Nistarika Nayar, Madhusoodanan & Molly genus novum 

Genus Leptochilo simillimum, sed etfoliis tri- vel quinquefariis in rhizomatedorsa liter 
dispositis et foliis fertilibus omnino sine lamina, sporangia vero nuda in costa 
gerentibus differt. Sorus linearis, exindusiatus, utroque costae latere continuus. 
Sporae monoletae, non perinatae. 
Typus: Nistarika bahupunctika Nayar, Madhusoodanan & Molly. 

A genus similar to Leptochilus but differing from it in having four rows of fronds 
restricted to the dorsal surface of the rhizome and in having the fertile frond totally 
devoid of any lamina but bearing sporangia directly on the midrib. Sorus linear, 
exindusiate and forming a lateral continuous line on either side of the midrib. 

Nistarika bahupunctika Nayar, Madhusoodanan & Molly sp. nov. 

Rhizoma ascendens, semi-epiphyticum, longum, sparse ramosum, plus minusve 
dorsiventraliter compressum, 5 - 8mm latum, 3 - 4mm crassum, viride, praeter ad 
apicem fere glabrum, dictyostelen faciens, sclerenchymate tenui fusoque in 
parenchymate dense disperse Paleae parvae, peltatae, clathratae, glandibus ad 
apicem et basi pilis duobus glanduliferis conspicuisque praeditae. Folia tri- vel 
quinquefaria dorsaliter in rhizomate irregulatim disposita, dimorpha, simplicia. Folia 
sterilia patentia, stipitata, lamina herbacea, angusto-lanceolata 30- 50cm longa(1.5)- 
2.5 - 4cm lata, in apicem acuminatum gradatim contracta, margine integro, venatione 
reticulata. Folia fertilia erecta, filiformia, sine lamina, 45 - 75cm longa 1.5mm lata. 
Sorus exindusiatus, linearis, continuus utroque costae nudae latere. Sporae 
monoletae, c. 35 x 65um, spinulosae, non perinatae. 



34 FERN GAZETTE VOLUME 13 PART 1 (1985) 



Holotypus: CU 29263, lectus ab K.K. Geeverghese et B.K. Nayar in 28.3.1980 apud 
Panthenthode, Silent Valley, India Meridionalis, et positus apud Central National 
Herbarium, Calcutta. 
Isotypus: positus apud Herbarium Universitatis de Calicut. 

Paratypi: positus apud Herbarium Universitatis de Calicut; CU 21329, lectus ab B.K. 
Nayar in 19.12.1977 et CU 29136 lectus ab B.K. Nayar et P.V. Madhusoodanan in 
13.2. 1980 apud Silent Valley. 

Rhizome climbing vertically up supports and becoming semiepiphytic, elongate, 
sparsely branched, dorsiventrally compressed, 5 - 8mm broad, 3 - 4mm thick, green 
nearly glabrous except near growing apices, dictyostelic and having slender, dark 
sclerenchyma strands profusely scattered in the ground tissue. Paleae small, peltate, 
clathrate, gland-tipped and bearing a pair of prominent glandular hairs at base. Fronds 
borne in four irregular rows on the dorsal surface of the rhizome, articulate to small 
phyllopodia, dimorphic and simple. Each frond of the two lateral rows is associated 
with a basal branch as well as a pair of roots, but the other fronds are not. Sterile fronds 
spreading (perpendicular to the rhizome), stipitate (stipe 1.5-1 2.0cm long, 1 .5 - 2.0cm 
thick, devoid of dorsal groove), with a thin, herbaceous, narrowly lanceolate lamina (30 
- 50cm long and (1 .5 -) 2.5 - 4.0cm broad) gradually tapered to a short acuminate apex 
and forming narrow tapered wings on the anterior region of the stipe; margin entire. 
Venation reticulate, having indistinct zigzag main veins and areoles possessing simple 
or forked included veinlets ending in hydathodes on the upper surface. Fertile fronds 
seasonal, erect (held parallel to the rhizome), clustered, filiform, devoid of lamina, 45 - 
75cm long and 1 .5mm broad. Sporangia aggregated in linear, exindusiate sori, one on 
either side of the naked midrib and extending all along it except for 5 - 1 2cm at the base 
and 1.0- 1.5cm at the apex, mixed with filamentous paraphyses;annulus 15 cells long. 
Spores rhonolete, c. 35 x 65pm, with spore coat not clearly differentiated into exine and 
perine, spinose. 

The name is derived from Sanskrit: Nistarika meaning lamina-less {ni = devoid of; 
starika = lamina) and bahupunctika meaning several rows [bahu - many; punctika - 
having rows). The generic n^me refers to the lamina-less nature of the fertile fronds 
and the specific name to the many rowed arrangement of fronds on the rhizome. 



MORPHOLOGY 
Rhizome 

N. bahupunctika grows in densely shaded forests among crowded undergrowth; it 
climbs up bases of woody shrubs and ultimately becomes semi-epiphytic as older 
parts of rhizome slough off. The rhizome is chlorophyllous, tenaciously attached by 
means of strong wiry roots borne in opposite pairs and horizontally encircling the 
support (Fig. 1 a); the posterior roots extend to the ground. The paleae are dense in the 
younger regions but sparse elsewhere and are peltate, blackish, prominently 
clathrate, subtriangular in shape (Fig. 2b), c. 1 -4 x 0.5mm, gland-tipped (Fig. 2c) and 
bearing a marginal uniseriate glandular hair (resembling the apex of the palea) on 
either side at the base. Ontogeny of the paleae is as in Leptochilus axillaris (Nayar 
1963). The rhizome is hard and brittle, with parenchymatous ground tissue having 
faintly thickened silvery walls bearing prominent pits; slender sclerenchyma strands 
(1-7 cells thick, having occluded lumen and thick blackish walls bearing prominent 
pits) are scattered throughout. The vascular cylinder is a perforated dictyostele having 
slender cylindrical meristeles, and is eccentrically placed in the rhizome (cortex 5 or 6 
cells thick on the ventral and 1 5-20 cells thick on the dorsal side). The endodermis is 
thin-walled but the inner walls of cells of ground tissue abutting on it are faintly 
thickened as in Paraleptachilus decurrens (not conspicuously thickened or dark 
brown as in L. axillaris). The dorsal half of the stelar cylinder is dissected by 4 or 5 
irregular longitudinal rows of overlapping frond gaps (Fig. 3a). The vascular 
connection to the frond is shallowly channel-shaped and dissected by three irregular 
longitudinal rows of lacunae. Each frond of the two lateral rows is associated with two 
roots but the median fronds do not have any root association. 

Study of apical organisation and morphogenesis in a large number of plants 
reveals that the patterns of growth and organogenesis in N. bahupunctika differ from 



NISTARIKA. A NEW GENUS OF POLYPODIACEAE 



35 





FIGURE 1 . Nistarika bahupunctika: a, a portion of the rhizome clinging to the substratum ana 
showing the clinging roots encircling the support; b, plant bearing a cluster of erect fertile fronds; 
(r, rhizome; c, clinging roots; f, fertile frond; d, posterior roots). 



36 FERN GAZETTE VOLUME 13 PART 1 (1985) 



those found in several related genera such as Drynaria, Microsorum, Paraleptochilus 
and Pyrrosia (Nayar and Molly unpublished). In these genera the rhizome apex forks 
into two after producing each frond, and a frond primordium is soon differentiated on 
each of the daughter apices on the side facing the sister apex. The daughter apex 
nearest the older frond becomes dominant and continues growth of the rhizome while 
the other one away from the frond (along with the frond primordium on it) becomes 
sluggish and ultimately dormant. The frond primordium on the dominant daughter 
apex grows out as a frond, pushing away to one side the dormant daughter apex which 
then appears as a dormant branch of the rhizome associated with the developed frond. 
Thus fronds are alternate and in two opposite rows on the rhizome and each is 
associated with an abaxial branch. In N. bahupunctika also the rhizome apex forks 
equally into two after producing a frond, and each daughter apex differentiates a frond 
primordium next to the apical meristem (Fig. 6a). However, in contrast to the taxa 
mentioned above, both frond primordia grow out so that a pair of closely placed fronds 
result (Fig. 6b). One of the daughter apices then becomes sluggish and ultimately 
dormant but a second frond primordium (opposite thef irst one) is established on it and 
this primordium remains dormant. The other daughter apex of the rhizome becomes 
dominant and soon forks into two, each of the resultant daughter apices bearing a 
frond primordium (Fig. 6c). Frond primordia on both daughter apices grow out as 
developed fronds so that a second set of two fronds is formed (Fig. 6d) one beside the 
other. The daughter apex away from the dormant apex of the earlier forking becomes 
sluggish and ultimately dormant while its sister apex (which is now medianly placed) 
becomes dominant, continuing growth of rhizome (Fig. 6e). It forks again and the 
process is repeated (Figs. 6f,g,h). The dormant apex of each forking gets pushed away 
to one side as the dominant sister apices continue to grow. This pattern of growth 
results in four (in some cases appearing as three to five) irregular longitudinal rows of 
fronds on the dorsal side of rhizome. Of these the two lateral rows are developed 
fronds borne on the dormant daughter apices of successive forking of the rhizome 
apex and only these fronds have a branch associated with each (the branch being the 
dormant daughter apex of each forking) at its abaxial base. The median rows of fronds 
are borne on the dominant daughter apices which have continued growth andthusdo 
not have any branch associated with them. 

Fronds 

The fronds are glabrous and glossy when mature; a few reduced paleae occur 
scattered on the stipe and main veins when young and a few minute (2-3 cells long) 
uniseriate non-glandular hairs on the lower surface of the lamina. Stomata are 
restricted to the lower epidermis and are copolomesoperigenous with the inner 
subsidiary cell narrow, having a smooth contour and nearly encircling the guard cells. 
The midrib is prominent and raised on both surfaces, but lateral veins are feeble and 
hidden in fresh material; the primary veins become somewhat evident in dried 
specimens. Primary lateral veins are 8-1 2mm apart and fork into divergent, strongly 
zigzag branches at the extreme base; the acroscopic branch is strongly oblique to the 
midrib and unites with the basiscopic basal branch of the next anterior vein to form a 
large costal areole, while the basiscopic branch is spreading, forking into equal 
divergent branches nearly 2 / 3 rds to margin. Basiscopic branches of successive main 
veins are interconnected by zigzag secondary veins to form two or three rows of 
primary areoles which are further subdivided into two or three secondary areoles, 
each of which has a divided included veinlet. There is a nearly regular row of small 
marginal areoles and beyond them an occasional free-ending veinlet pointing to the 
margin. The large costal areoles possess a solitary included veinlet pointing to the 
midrib and commonly forked with the branches strongly divergent. All vein endings are 



NISTARIKA, A NEW GENUS OF POLYPODIACEAE 



37 



clavate and end in hydathodes in the upper epidermis. Fertilefronds are produced only 
during the dry season; they are usually 2-4 (up to 8) per apex of rhizome, held nearly 
parallel to the rhizome and extend a long way anterior tothe rhizome apex. A lamina is 
totally absent and sporangia are borne directly on the midrib, with the sorusforming a 
continuous line on either side at the position where a lamina should have been. At the 
region of the sorus the midrib is grooved, with the soral placentum occupying the base 
of the groove (Fig. 4c,d). 

Sporangia, spores and prothalli 

The sporangia are mixed with slender, uniseriate, multicellular (5-6 cells), elongated, 
filamentous, gland-tipped paraphyses (Fig. 4d). The sporangial stalk is slender and 
elongated (5 cells long); annulus is 15 (14-16) cells long and prominently indurated. 
The spores (Fig. 2a) are bilateral, concavo-convex in lateral view, elliptic-oblong in 
polar view and 35 x 65um in size but swelling to 40 x 75pm when acetolysed. The 
spore coat is golden-brown, less than 1.5um thick and densely spinose; spines 
irregularly aggregated in groups, 5-7pm long, less than lumthick, sometimes moreor 
less curved, blunt and somewhat deciduous (mostly shed on acetolysis). The adult 
prothallus is of the ribbon-like branched type (Nayar and Kaur 1970) as \r\Leptochilus, 
forming small clustered patches. It is naked and one cell thick except for small 
cushions 2-4 cells thick borne in an irregular row medianly on the ribbon-like 
branches. Sex organs are borne on the lower surface of these cushions and are of the 
common type in Polypodiaceae. 




FIGURE 2. Nistanka bahupunctika: a, lateral view of spore; b, mature pa lea, c, apex of mature 
palea; (s, stalk of palea). 



38 



FERN GAZETTE VOLUME 13 PART 1 (1985) 




FIGURE 3. a. Dorsal view of vascular cylinder of a portion of rhizome; b, ventral view of vascular 
cylinder of the same portion of rhizome (b, branch trace; c, branch gap; g, frond gap; I, frond trace; 
m, medium ventral row of lacunae; r, root trace). 

DISCUSSION 
In general, Polypodiaceae sensu stricto are characterised by having a creeping 
dorsiventral rhizome bearing two dorsal rows of fronds (in some species, e.g. 
Microsorum linguaeforme and Pseudodrynaria coronans, the rows are so close as to 
appear as one), each articulated to a phyllopodium and associated with an abaxially 
lateral branch bud. Oleandropsis Copel. is exceptional in having an erect-growing 
slender radially symmetrical rhizome bearing several rows of fronds all around it. 
Nistarika differs from all other Polypodiaceous genera in having four rows of fronds 
restricted to the dorsal surface of a dorsiventral rhizome, and having branch buds 
associated with only the two lateral rows of fronds, the other fronds having no branch 
association. The growth habit of the plant which results in this characteristic 
arrangement of fronds and branches is unique, but only a variation from the common 
pattern found in the family (apical forking followed by one of the resultant branches 
becoming dormant). Sporangia are borne on the lamina of fertile fronds in all known 
Polypodiaceae; the lamina is highly reduced in some, but sporangia are restricted to 
the lamina region only and the midrib does not bear any sporangia. In contrast there is 
no trace of lamina in the fertile fronds of Nistarika; sporangia are borne directly on the 
midrib, and the fertile region forms a groove on either side, with the soral placentum 
seated at the base of the groove. However, the vascular supply to the sorus resembles 
the main lateral veins of the sterile lamina in its origin from the vacular bundles of the 
midrib, but remains well within the cortex of the midrib. Though a lamina is absent, the 
linear sori occupy the position that would be expected, were a lamina present. The 
combination of these unique characters excludes Nistarika from any known genus of 
ferns, justifying its separation as a new genus. 



NISTARIKA, A NEW GENUS OF POLYPODIACEAE 



39 




10 mm 

FIGURE 4. Nistarika bahupunctika: a, venation pattern, b, transection of a very young fertile frond 
showing initiation of sori; c, transection of a mature fertile frond; d, transection of sorus. 
(a, margin of frond; f, soral vascular supply; I. lateral vein subtending sorus; m, midrib; p. 
paraphysis; s, sporangium; t, third row of stalk cells; v, vascular bundle of midrib). 

Nistarika resembles Leptochilus Kaulf. and Paraleptochilus Copel. except for the 
characteristic peculiarities of fertile frond, multi-rowed arrangement of fronds and 
eccentrically placed vascular bundle of rhizome. Though dried herbarium material of 
N. bahupunctika is confusingly similar in appearance to dried material of/., axillaris 
and P. decurrens forma decurrens and forma lanceolata, the resemblance is not so 
close with fresh material in the field. The habit of the four genera is characteristically 
different (Table 1 )and/V. bahupunctika differs from the others in its fertile fronds being 
clustered in groups and held vertically (paralleltothe verticalsubstratumonwhich the 
plant grows, in contrast to being at c. 90° with the substratum in others); their lamina- 
less nature renders them markedly more slender than the fertile fronds of other taxa. A 
comparison between N. bahupunctika and L. axillaris, P. decurrens forma decurrens 
and forma lanceolata is given in Table 1 ; Figure 5 gives a comparison of the venation 
pattern of the sterile frond. The venation of P. decurrens forma lanceolata is quite 
similar to N. bahupunctika, but that of P. decurrens forma decurrens is clearly 
different, particularly the very prominent nature of the primary lateral veins in the 
latter, and in the possession of a sheath of cells with coloured (dark-brown to blackish) 
deposits, making them more prominent. In possessing prominent sclerenchyma 
strands in the rhizome, and s'ender meristeles devoid of a prominent thick walled 
sheath, N. bahupunctika resembles P. decurrens; venation of sterile fronds and 
possession of filamentous paraphyses are additional similarities with P. decurrens 
forma lanceolata, but the semi-epiphytic habit, lack of sclerenchyma strands in the 
ground tissue of the stipe, possession of a thick-walled sheath around the vascular 
bundles of the stipe, the peltate nature of paleae, and spores bearing bacula-like large 
spines distinguish N. bahupunctika from P. decurrens. Though N. bahupunctika 
resembles L. axillaris in possessing similar paleae, absence of sclerenchyma strands 
in the stipe, possession of a thick walled sheath around the vascular bundles of the 
stipe, the venation pattern of the sterile leaf and possession of filamentous 



40 



FERN GAZETTE VOLUME 13 PART 1 (1985) 



paraphyses, it differs in possessing profuse sclerenchyma strands in the rhizome, 
lacking a thick walled sheath around meristeles, and having a stout and comparatively 
short rhizome which is tenaciously attached to the substratum. In the ribbon-like 
nature of its prothallus N. bahupunctika resembles both Leptochilus and 
Paraleptochilus, but it differs from both these genera in the pattern of its growth, 
lamina-less fertile fronds and bacula-like large spines on the spore wall. 

TABLE 1. Comparative morphology of Nistarika, Leptochilus and Paraleptochilus 



Sporps 



Paraphyses 



Nistarika bahupunctika 

Restricted to densely shaded 
forest beds; initially terre- 
strial (creeping on soil) but 
later climbing up bases of 
supports and ultimately semi- 
epiphytic with rhizome not in 
contact with soil but bearing 
many roots extending to soil 
below. 

Creeping vertically up, tena- 
ciously attached to substratum 
ca. 8 mm thick, moderately 
elongated, hard, brittle, 
sparsely bianched and having 
profuse sclerenchyma strands 
in ground tissue. 

Peltate deciduous (absent in 
older regions). 

Dictyostelic with many broad 
rather short perforations; 
meristeles slender, devoid of 
conspicuous sheath. Stelar 
cylinder eccentrically placed 
in the rhizome. 



In 4 dorsal rows and only 
lateral ones associated with 
a branch each; stipe devoid 
ot sclerenchyma strands but 
vascular bundles possessing a 
thick walled sheath. Fertile 
leaves clustered, devoid of 
lamina and bearing spoiangia 
directly on midrib. 



Bilateral, 35 x 65 aj, swelling 
to 40 x 70 Li on acetolysis; 
wall spinose with spines 7 ti 
long, blunt and uniformely 
thick throughout. 



Filamentous, elongated, many 
cells long. 



Leptochilus axillaris 

Epiphytic in partially 
shaded areas, attached 
to tree trunks, growing 
throughout life with no 
connection to soil and 
wide creeping. 



Creeping horizontally, 
loosely attached to 
substratum, ca. 5 mm 
thick, extensively 
elongated, soft, pro- 
fusely branched, devoid 
of sclerenchyma. 

Peltate, deciduous (ab- 
sent in older regions). 

Dictyostelic with very 
narrow, much elongated 
perforations; meristeles 
thick, having a promi- 
nently conspicuous 
dark sheath of thick 
wallea cells Stelar 
cylinder not eccentrically 
placed in rhizome. 
In 2 dorsal rows and each 
associated with a brand i, 
stipe devoid of sclerenchyma 
strands but vascular bundles 
possessing thick-walled 
sheath. Fertile leaves not 
clustered, having very 
narrow lamina bearing 
spoiangia; midrib devoid 
of sporangia. 

Bilateral, 30 x 54 y 
not swelling on aceto- 
lysis; wall spinulose 
with spinules 3 n long, 
tapered to sharp apex. 

Filamentous, elongated, 
many cells long. 



Paraleptochilus decurrens P decurrens forma lanceolate 



Terrestrial in densely 
shaded forest beds near 
streams and waterways; 
short creeping on rocky 
substrata. 



Creeping horizontally, 
tenaciously attached to 
substratum, ca. 8 mm thick 
more or less short, sparsely 
branched, brittle, hard, 
having profuse sclerenchyma 
strands in ground tissue. 

Basally attached, dense all 
over, non-deciduos. 



Dictyostelic with many 
broad, rather short perfo- 
rations; meristeles slender, 
devoid of conspicuous 
sheath. Stele not eccenti i- 
cally placed in rhizome. 



Terrestrial in partially 
shaded forest beds; wide 
creeping on rocky or gravally 
soil but with a tendancy to 
climb up supports. 



Creeping horizontally, loosely 
attached to substratum ca. 
6 mm thick, much elongated, 
hard brittle, profusely 
branched and having piofuse 
sclerenchyma strands in 
ground tissue. 

Basally attached, non- 
deciduous, but not dense in 
older regions. 

Dictyostelic with many broad, 
rather short perforations; 
meristeles slender devoid of 
conspicuous sheath. Stele not 
eccentrically placed in 
rhizome. 



In 2 dorsal rows and each 
associated with a branch; 
stipe having slender scle- 
renchyma strands in the 
ground tissue but vascular 
bundles devoid ot thick- 
walled sheath. Fei tile leaves 
not clustered, having very 
nanow lamina beating 
sporangia, midrib devoid 
of sporangia. 

Bilateral, 37 x 55 u, not 
swelling on acetolysis; wall 
spinulose with spinules 3 u 
long, slender, tapered to a 
sharp apex, deciduous, irre- 
gularly clustered. 

Absent. 



In 2 dorsal rows and each 
associated with a branch; 
stipe having slender scleren- 
chyma strands in ground tissue 
but vascular bundles devoid of 
thick walled sheath. Fertile 
leaves not clustered having 
very narrow lamina bearing 
sporangia; midrib devoid of 
sporangia. 

Bilateral, 35 x 57 u, not 
swelling on acetolysis; wall 
spinulose with spinules 5 jj 
long, slender, tapered to 
a sharp apex, aggregated in 
irregular groups. 

Filamentous, elongated, many 
cells long. 



N. bahupunctika resembles some species of Bolbitis Schott in its characteristic 
frond arrangement and association of branch buds with frond bases (Nayar and Kaur 
1965). In addition several species of Bolbitis possess a highly reticulate venation 
similar to N. bahupunctika and a highly reduced fertile frond; also, at least some 
species of Bolbitis possess simple fronds. A tendency for the rhizome to climb vertically 
up bases of woody shrubs and for the plant to become semi-epiphytic like N. 
bahupunctika is seen in B. appendiculata (several such plants were noted, growing 
side by side with N. bahupunctika in Silent Valley), and a similar habit is reported for B. 
sinensis in Thailand (Hennipman 1977). In this context it is interesting to note that 
Hennipman (1977) in his monographic study of Bolbitis reported that Leptochilus 
trifidus v.A.v.R. of Sumatra is a Bolbitis and possibly a hybrid with Leptochilus, thereby 
suggesting some degree of relationship between the two genera. However, Bolbitis 
differs from N. bahupunctika as well as Leptochilus in its non-peltate paleae, non- 
articulated fronds, the stelar cylinder of the rhizome having an intact broad basal half, 
possession of foliar bulbils and chromosome number (n = 41 , in contrast to n = 36, 37 in 
Polypodiaceae). Even the suspected hybrid, L. trifidus, possesses basally attached 



NISTARIKA, A NEW GENUS OF POLYPODIACEAE 



41 





FIGURE 5. Venation of sterile frond of a, Nistarika bahupunctika, b, Paraleptochilus decurrens 
forma lanceolata and c, P. decurrens var. decurrens. 




FIGURE 6. Diagrammatic representation of pattern of growth in Nistarika bahupunctika (for 
details see p. 36). 



4? FERN GAZETTE VOLUME 13 PART 1 (1985) 



pseudo-peltate paleae bearing stout glandular marginal hairs, non-articulated leaves, 
foliar bulbils and a vascular cylinder having a prominent ventral meristele and 
prominently perinate spores, all characteristic of Bolbitis. 

Nistarika is evidently related to Leptochilus and like it derived from Microsorum. 
Its epiphytic habit, elongated dorsiventral rhizome climbing vertically up supports and 
bearing stout long roots reaching down to the soil, simple lanceolate horizontally 
spreading leaves, reticulate venation with zigzag inconspicuous main veins which 
extend only part way up to the margin, filamentous paraphyses, 1 4-1 6 celled annulus 
and the large bilateral spores suggest M. superficiale (Bl.) Ching, but in the extreme 
reduction of the fertile lamina it is more advanced than Microsorium and even 
Leptochilus. 

ACKNOWLEDGEMENTS 
We should like to thank K. Kavanagh for correcting the Latin description. 

REFERENCES 
HENNIPMAN, E. 1977. A monographic of the fern genus Bolbitis (Lomariopsidaceae). Leiden 

University Press. 
NAYAR, B.K. 1 963. Contribution to the morphology of Leptochilus and Paraleptochilus. Amer. J. 

' Bot. 50: 301 -308. 
NAYAR, B.K. 1 982. Flora and Fauna of Silent Valley. Attappadi and Sabarigin Forests. 

K.S.E. Board, Trivandrum. 
NAYAR, B.K. and S. KAUR. 1965. Studies on the fern genera Bolbitis and Egenolfia — I: 

Morphology of the sporophytes. J. Linn. Soc. (Bot.) 59: 1 27-1 40. 
NAYAR, B.K. and S. KAUR. 1970. The yametophytes of homosporous ferns. Bot. Rev. 37: 295- 

396. 

SHORT NOTE 

CYRTOMIUM FALCATUM IN IRELAND 

A single plant, identified as Cyrtomium fa/catum(L.) Presl was located in west Cork, on 
the south coast of Ireland in December 1983. This is the first record of this species 
naturalized in Ireland. The plant grew in the seaspray zone of a 3m high cliff face with a 
public pathway just overhead, and had fourteen mature fronds, five young fronds and 
six dead ones. One dead frond measured 82cm. Young fronds were light green in 
colour and soft whereas the older ones were darker green and more cartilaginous. The 
brown scales were up to 1.3cm long andthe pinnae up to 8cm long. The fronds tapered 
towards the top and less abruptly towards the bottom and were devoid of pinnae on the 
lower half. The pinnae had very distinct main veins with the numerous small orbicular 
sori scattered in between the secondary veins. Sori nearer the margins were closer 
together and smaller than those nearer the main vein. The indusium was circular and 
visible even in December. Of the five young circinately coiled fronds observed in 
December three were fully expanded four weeks later and the rudimentary sori were 
even faintly visible at that stage. Some pinnae while otherwise very healthy looking 
appeared to have been grazed (cf. specimen lodged in DBN), while on other fronds the 
pinnae were distorted and convoluted possibly as the result of seaspray or a virus or 
fungal infection. 

It appears to be naturalized in this seaside location in west Cork. Likewise the 
species has been found in Australia colonising coastal cliffs and old cuttings near the 
sea and under boulders on the shore in the Isles of Scilly. The origin of this Irish 
specimen is unknown. The nearest iikely source is a house a half mile away. 

J.P. CULLINANE & C. CROWLEY 
Department of Botany, University College, Cork, Ireland 



FERN GAZ. 13(1) 1985 43 



THE GENERIC IDENTITY OF POLYPODIUM BANAENSE 



W.LA. HETTERSCHEID 

Institute of Systematic Botany, State University of Utrecht, Heidelberglaan 2, 
P.O. Box 80.102, 3508TC, Utrecht, The Netherlands 

ABSTRACT 

The species Polypodium banaense C.Chr. is transferred to Crypsinus. The 
recognition of a genus Phymatopteris Pic. Ser. (= Phymatops/s J. Sm.) separate from 
Crypsinus is discussed. 

INTRODUCTION 
In the course of my studies on the venation of Polypodiaceae, I visited the Paris 
herbarium and came across a few sheets of the indochinesefern originally published 
by Christensen (1934a) as Polypodium banaense. Subsequent character analysis of 
the venation, frond shape, rhizome scales and spores supports the view that the 
species belongs to Crypsinus (Polypodiaceae s.str.) as interpreted by Copeland (1 947), 
and is therefore referred to that genus. 
Crypsinus banaensis (C.Chr.) Hetterscheid comb. nov. 

Polypodium banaense C.Chr., Bull. Mus. (Hist. Nat., Paris), 2e se>, 1934: 105. 
Phymatodes banaense (C.Chr.) C.Chr. et Tard., Not. Syst., 8(4), 1939: 190. Tard. & 
C.Chr. in Lecomte; Fl. gen. Indoch., 7(2) fasc. 9, 1941: 468. Paragramma banaensis 
(C.Chr.) Ching, Sunyatsenia 5(4), 1940: 258. Type: Sallet (Herb. Ecole sup. d'Agric. 
Hanoi, n. 3534), (BM, non vidi). 

TAXONOMIC HISTORY 
In the same year of its publication, Christensen listed Polypodium banaense in his 
Index Filicum (1934b, suppl. tert.) as belonging to Polypodium subg. Microsorium. 
Christensen & Tardieu-Blot (1939) transferred the species to Phymatodes Presl (= 
Phymatosorus Pic. Ser.) sect. Paragramma, on account of its indistinct venation, as 
opposed to sect. Euphymatodes. Ching (1 940) accepted Paragramma Moore as a valid 
genus and proposed Paragramma banaensis (C.Chr.) Ching, which was not followed 
by Tardieu-Blot & Christensen (1941), who retained the species in Phymatodes, still 
regarding sect. Paragramma as part of it. 

OBSERVATIONS 
In addition to the descriptions given by Christensen (1934a) and Tardieu-Blot & 
Christensen (1 941 ) the following observations are considered to be relevant. Contrary 
to Christensen's (1934a) remarks, the species is not dimorphic. Gradual frond 
elongation in both sterile and fertile specimens exists. The margin of small fronds is 
regularly notched (Fig. 1a, b), whereas this feature becomes irregular in fronds of 
intermediate lengths (Fig. 1c), and is absent in the largest fronds (Fig. 1d,e). 

The venation (Fig. 1 ) consists essentially of the following parts: a series of narrow 
elongated areoles on either side of the rhachis, in small fronds empty (Fig. 1a, b), in 
larger fronds sometimes containing one free recurrent vein (Fig. 1d,e). Each of these 
rows is bordered by a row of large more isodiametrical areoles. The included venation 
of the latter in the smallest fronds consists of one or two, usually free, excurrent veins 
(Fig. 1a, b). In all larger fronds the included venation is branched and anastomosed, 
thus developing smaller types of areoles (Fig. 1c,d,e), which are usually empty. Outside 
these larger areoles excurrent veins exist, which sometimes anastomose to form small 
marginal areoles. 



44 



FERN GAZETTE: VOLUME 13 PART 1 (1985) 




FIGURE 1. Venation patterns in Crypsinus banaensis (C.Chr.) Hetterscheid. a,b, small sterile 
fronds; c, intermediate sterile frond; d, large sterile frond; e, large fertile frond. (a,b, Sallet s.n. (P); 
c, Poilane 3524 (P); d,e, Poilane 51 1 1 (P).). Scale bar: 5 mm. 



THE GENERIC IDENTIFY OF POLYPODIUM BANAENSE 45 



The sori are situated in the larger areoles on fusion points in the included 
venation, one in each of them (Fig. 1e). 

The spores are of the Microsorium-Xype (Hennipman & Roos 1983) and have a 
thick perispore as found in all species of Crypsinus and in some of thedrynarioidferns 
(pers. comm. E. Hennipman). 

Contrary to Christensen (1934a) and Tardieu-Blot & Christensen (1941), I was 
unable to find receptacular paraphyses. The filiform paraphyses referred to are 
probably decapitated sporangial stalks which may be abundant in the sori. 

DISCUSSION 
A number of arguments indicate that inclusion of Polypodium banaense in either 
Phymatosorus or Paragramma is not justified. The venation is in essence different 
from that in both genera, the latter superficially resembling a goniophlebioid pattern, 
but possessing a much more complicated included venation in the larger areoles, and 
developing from a very different series of blastogenetic stages (Mitsuta 1981, 
Hetterscheid & Hennipman 1984). The rhizome scales in both genera are clathrate, 
whereas those in P. banaense are opaque. The spores of P. banaense do not 
correspond to the type found in Paragramma or typical Phymatosorus, which possess 
the rather unique lepisorioid type (Hennipman & Roos 1 983). Furthermore species of 
Paragramma possess peltate paraphyses of the type found in Lepisorus, which are 
absent in the present species. 

The venation of the young fronds, the rhizome scales, the notching of the small 
fronds, and the spores all clearly correspond to features found in species of Crypsinus. 
Christensen (1934a) mentioned the strong similarity of P. banaense to P. 
stenophyllum Bl. (= Crypsinus stenophyllus (Bl.) Holtt.). He also compared the present 
species to P. rhynchophyllum Hook. (= Phymatopteris (Crypsinus) rhynchophylla 
(Hook.) Pic. Ser.). 

The distinctions given by Ching (1 964) to separate Phymatopteris from Crypsinus 
s.str., do not seem to be very conclusive. The species Polypodium (Crypsinus) 
ensiforme Thunb.* for instance, has a very striking goniophlebioid venation (a 
character used by Ching to delineate Crypsinus s.str.), whereas the distinct lateral 
veins and large sori and pinnatifid frond point to an inclusion in Phymatopteris. 
Furthermore, in narrow frond parts of certain species of Phymatopteris, the venation 
reduces to a goniophlebioid pattern (e.g. P. albidosquamata (Bl.) Pic. Ser.; Mitsuta 
1984, Figs. 580-581). The "drynarioid" venation mentioned as a character of 
Phymatopteris is not useful, as a number of species of Drynaria have a goniophlebioid 
venation {e.g.D.parishii, D. sinica; pers. comm. M.C. Roos). I therefore suggest that the 
genus Crypsinus be used in its broadest sense (Copeland 1947) until it is dealt with 
monographically. 

The venation of Crypsinus banaensis shows an intriguing mixture of characters 
common to both Crypsinus and Microgramma Presl/ Pleopeltis Humb. et Bonpl. ex 
Willd. The small fronds contain a venation found in many young specimens of 
Crypsinus species or allies (e.g. Pycnoloma C.Chr.). the adult venation on the other 
hand shows a striking similarity to that found in species of Microgramma (compare Fig. 
lewithdelaSota &Pe>ez-Garda 1982, Fig. 3a,b), or P/eope/f/s (e.g. Mitsuta 1981, Fig. 
6, 11, 17). In the light of morphogenesis of adult venations in blastogenetic frond 
series this is an interesting observation, showing that the Microgramma type of 
venation can develop via two different pathways, one, in most species of 
Microgramma, following the pattern found in Polypodium L. and Goniophlebium Presl, 
and another following part of the development in most Crypsinus species (Mitsuta 
1984). 

*The suggestion that Phymatodes ensiformis belongs to Crypsinus is debatable (it certainly is not 
a Phymatodes). 



46 FERN GAZETTE: VOLUME 13 PART 1 (1985) 



MATERIAL STUDIED 
Poilane 3524 (P), 51 1 1 (P), 6925 (P), 23913 (P); Sallet s.n. (P). 

ACKNOWLEDGEMENTS 
Thanks are due to Prof. Dr E. Hennipman for stimulating the preparation of the 
manuscript as well as its correction, and for providing information on the spores, and 
to Miss G.P. Verduyn and Mr M.C. Roos for additional information. 

REFERENCES 
CHING, RC. 1940. On natural classification of the family "Polypodiaceae". Sunyatsenia 5: 201- 

' 268. 
CHING, RC. 1 964. On the genera Phymatopsis J.Sm. and Crypsinus Presl. Acta Phytotax. Sinica 

'9: 179-197. 
CHRISTENSEN, C. 1 934a. Filices novae indochinensis.flty//. Mus. (Hist. Nat, Paris), 2es6r 6: 100- 

106. 
CHRISTENSEN, C. 1934b. Index Filicum. Supplementum tertium pro annis 1917-1933. 
CHRISTENSEN, C. & TARDIEU-BLOT, M.-L 1939. Les fougeres d'lndochine: XV. Dipteriodeae- 

XVI. Polypodioideae - XVII Elaphoglossoideae. Not. Syst. 8: 175-210. 
COPELAND, E.B. 1947. Genera Filicum. Waltham, Mass., Chronica Botanica. 
HENNIPMAN, E. & ROOS, M.C. 1983. Phylogeneticsystematicsof the Polypodiaceae (Filicales). 

Verh. naturwiss. Ver. Hamburg (NF) 26: 321-342. 
HETTERSCHEID, W.L.A. & HENNIPMAN, E. 1984. Venation patterns, soral characteristics, and 

shape of the fronds of the microsorioid Polypodiaceae. Bot. Jahrb. Syst. 105: 1 1-47. 
MITSUTA, S. 1981. Venation of Lepisorus and Pleopeltis (Polypodiaceae). Acta Phytotax. Geobot. 

32: 147-164. 
MITSUTA, S. 1984. Studies in the venation and systematics of Polypodiaceae. II. Venation of 

Polypodiaceae (2) and of some other ferns. Mem. Fac. Sc. Kyoto Univ., ser. biol. 9: 57-85. 
SOTA, E.R. DE LA & PElREZ-GARCIA, B. 1982. Nervacion y dimorfismo foliar en Microgramma 

Presl (Polypodiaceae s.str.). Biotica 7: 45-64. 
TARDIEU-BLOT, M.-L. & CHRISTENSEN, C. 1941. "Fougeres". In H. Lecomte, Flore g6n6rale de 

I'lndo-Chine 7(2) fasc. 9 (1941): 433-544. 



REVIEW 

ARKANSAS FERNS AND FERN ALLIES by W. Carl Taylor, illustrated by Paul 
W. Nelson. 262 pp. 240 x 207mm. Published by the Milwaukee Public 
Museum ISBN 0-89326-097-5 Arkansas. 1984. Price $29.00. 

This is a first-rate, beautifully produced book, the contents and presentation of which 
are an example to all local flora writers. I use the phrase 'local flora' with hesitation; 
Arkansas is larger than the whole of England by 3000 sq. miles! Introductory chapters, 
brief yet to the point, explain ferns and allied plants to the naturalist and layman; an 
illustrated glossary is a plus; pteridophyte distribution and ecology, discussed 
naturally in an Arkansas context, make interesting reading. 

A well illustrated key is given to the 32 genera included. The bulk of the book is 
given to descriptions of these genera, keys to the 78 species found in Arkansas, and 
clear descriptions with a full page line-drawing of each species by one of the leading 
botanical artists of the present time. Not only is each page beatifully laid out but the 
close-up drawings are accurate and show what is required. As so many of the species 
are either grown in Britain or related to the European flora I expect the book to be as 
useful on this sides of the Atlantic as on the other. 

A. C. JERMY 



FERN GAZ. 13(1) 1985 47 



THE PTERIDOPHYTE HERBARIUM OF 
TRINITY COLLEGE DUBLIN 



J. PARNELL 
School of Botany, Trinity College, University of Dublin, Dublin 2, Ireland 

ABSTRACT 

The pteridophytes in the herbarium of Trinity College Dublin (TCD) are described. 
They comprise about 8000 specimens from 417 collectors and include previously 
unrecognised isotypes and holotypes. A list of major collectors is given with 
localities. 

INTRODUCTION 
The herbarium of Trinity College Dublin (TCD) was established around 1834, is the 
second largest herbarium in Ireland and contains at least 200,000 specimens 
(including lichens and bryophytes) with a notable collection of flowering plants and 
algae (see Holmgren, Keuken and Schofield, 1981). The only published list of 
collectors is that of Parnell (1982a) which relates solely to the rather small lichen 
collection. The pteridophyte collection is completely undocumented. 

THE PTERIDOPHYTE COLLECTION AT TCD 
Until recently the pteridophyte collection was ordered according to the scheme of 
Copeland (1 947). Over the past two and a half years the collection has been re-ordered, 
to correspond with the scheme detailed by Crabbe, Jermy and Mickel (1975); at the 
same time nomenclature was revised to follow the most recent appropriate literature. 

The collection totals approximately 8000 specimens, representing 228 genera 
collected largely during the 1 9th and early 20th centuries, with many specimens from 
remote and rarely visited localities, for example New Caledonia, St Helena and Oahu in 
the Hawaiian Islands. Perhaps the most interesting small collection is that made by 
W.J. Sollas on a famous Royal Society expedition in 1 896, which made deep borings 
into the coral of Funafuti, a small island in the SW Pacific near the Ellice Islands, 
thereby confirming Darwin's theories on the origins of coral reefs. 

Approximately one eighth of the specimens in TCD come from the British Isles 
and, of these, one sixth were collected by the previous curator Prof. D.A. Webb. The 
European collection (including the British Isles) totals some 1400 specimens and is the 
one most added to in the latter part of this century. The collection also has a large 
number of specimens from Australasia, India, Sri Lanka and tropical South America. 

Revision has indicated that the collection contains a number of isotypes and some 
holotypes. The majority of the 417 collectors among the pteridophytes are also 
represented in the phanerogamic collection. The most prolific and important collectors 
are listed below; countries of origin are given after the collector. South Africa is taken 
to exclude South West Africa (Namibia) but to include Lesotho, Swaziland and all 
Bantustans. 

Archer, W. (Tasmania); Babington, C.C. (China and Japan); Backhouse, J. (Mauritius); Ball, J. 
(Algeria, France, Germany, Italy, Morocco, Sicily, Switzerland, Spain); Ball, P.W. StChater, A.O. 
(Spain, Jugoslavia); Barnard, J.C. (Trinidad); Barter, C. (Fernando Po, Sierra Leone); Beckett, 
T.W.N. (Sri Lanka); Brown, W. (China, west coast of Africa); Buckley, S.B. (U.S.A.); Claussen, P. 
(Brazil); Cooper, T. (South Africa); Coulter, T. (Britain, Ireland, Madeira, Mexico, Panama, U.S.A.); 
Cuming, H. (Chile, Galapagos Islands, Malacca, Panama, Philippines, Pitcairn Island, Society 
Islands, St Helena); Deplanche, ? (New Caledonia); Drege, C.F. (South Africa); Drummond, T. 
(Canada, U.S.A.); Eaton, D.C. (Filices Boreali-Americanae); Edwards, Rev. ? (Jamaica); Fendler, A. 
(New Mexico, Panama, Venezuela); Gardner, G. (Brazil, India, Sri Lanka); Gerrard, W.T. (South 
Africa); Gerrard, W.T. & McKen, M.J. (South Africa); Griffith, W. (China, India, Malaysia, Malesia, 



48 FERN GAZETTE: VOLUME 13 PART 1 (1985) 



Philippines, Sri Lanka); Gunn, R.G. (Tasmania); Halliday, G. (Britain, Ireland); Harvey, W.H. 
(Australia, Chile, Fiji, Friendly Islands, Great Britain, Ireland, Madeira, New Zealand, Sri Lanka, 
South Africa, Switzerland, Tasmania, U.S.A.); Haughton, ? (St Helena); Hind, W.M. (Britain, 
Ireland); Hooker, J.D. (Argentina, Australia, Brazil, Guyana, Canada, Columbia, Falkland Islands, 
Guatamala, Hawaii, Hermite Island, India, Jamaica, Java, Kerguelen Island, Madeira, Mauritius, 
Norfolk Island, Oahu, Peru, St Helena, Sandwich Islands, Sierra Leone, South Africa, Sri Lanka, 
Switzerland, Tasmania, Trinidad); Hooker, J.D. & Thomson, T. (Bangladesh, India); Hooker, W.J. 
(ex. herb. Jamaica, Madagascar, Madeira, Mauritius, Sri Lanka); Hunt, T.C. (Azores); Hutton, H. 
(South Africa); Jameson, W. (Brazil, Ecuador, Mexico); Jenkins, F. (India); Johnson, E. (India); 
Kelly, D.L (Jamaica, Ireland); Kerr, A.F.G. (Siam); Kirk, T. (ex. herb. Britain); Lechler, W. (Chile, 
p eru _ a set issued by Hohenacker); Linden, J.J. (Nicaragua, Venezuala); Lobb, T. (Java, Malaya); 
Lyall, D. (Australia, Greenland, New Zealand, South Sea Islands); Mackay, T.J. (Ireland); 
Mathews, A. (Peru, Society Islands); Miers, J. (Brazil); Oldham, T. (India); Oakes, W. (U.S.A.); 
Pearson, C. (Ireland); Poeppig, E.F. (Brazil, Chile, Cuba, Peru, U.S.A.); Pringle, A. (Ireland); 
Sartwell, H.P. (U.S.A.); Schomburgk, R.H. (Guyana); Schultz, N. (Norfolk Island); Simons, ?(lndia); 
Sinclair, A. (some labelled ex. herb. W. Gourlie, New Zealand, Mexico, Panama); Spruce, R. 
(Brazil, Ecuador, Peru); Strange, F. (Australia); Thomson, T. (India); Thwaites, G.H.K. (Sri Lanka); 
Vieillard, E. (New Caledonia); Wallich, N. (India, Malaysia and some enigmatically labelled East 
Indies (see Van Steenis [1950-]); Webb, D.A. (Australia, Austria, Britain, Bulgaria, France, 
Ireland, Italy, Jugoslavia, Madeira, Portugal, Spain, Sweden, Switzerland, Turkey); Wright, ? Miss 
(the daughter of C.H. Wright) (Britain); Wright, C. (Cuba, New Mexico, South Africa); Wright, E.P. 
(Ireland). 

The pteridophyte collection is now in the process of computerisation (Parnell 
1982b), a process which it is hoped will eventually be extended to the whole 
herbarium. Further details of the collection, its contents and additional information on 
collectors are available on request 

ACKNOWLEDGEMENTS 
I would like to thank Miss E. Lowe for extracting some of the data relating to collectors 
f r om the herbarium, Dr D.L. Kelly and Professor D.H.S. Richardson for helpful 
comments on an early draft of this manuscript. 

REFERENCES 
COPELAND, E.B. 1947. Genera filicum. The genera of ferns. Chronica Botanica, Waltham, 

Massachusetts, U.S.A. 
CRABBE, J. A., JERMY, A.C. & MICKEL, J.T. 1 975. A new generic sequence for the pteridophyte 

herbarium. Fern Gaz. 11: 141-162. 
HOLMGREN, P.K., KEUKEN, W. & SCHOFIELD, E.K. 1981. Index Herbariorum. Part 1 The 

Herbaria of the World. Ed. 7. Dr W. Junk B.V. The Hague/Boston. 
PARNELL, J.A.N. 1 982a. The lichen herbarium of Trinity College Dublin (TCD). Lichenologist 14: 

280-281. 
PARNELL, J.A.N. 1982b. The use of the herbarium in biological mapping and recording. 

Environmental Education Newsletter 19: 4-7. 
VAN STEENIS, C. G.G.J. 1950-. Flora Malesiana. Noordoff-Kolff, Djakarta. 



FERN GAZ. 13(1) 1985 49 



SPOROPHYLL-PTERYX' IN AFRICAN AND AMERICAN 

SE LAG IN ELLA 



NAT. QUANSAH and BARRY A. THOMAS* 

Life Sciences Department, University of London, Goldsmiths' College, 

Rachel McMillan Building, Creek Road, London, SE8 3BU, England 

ABSTRACT 

Vertical /oblique projections are reported as occurring on the adaxial surfaces of 
sporophylls of fourteen West African and twenty South American species of 
Selaginella. We propose that this projection be called 'sporophyll-pteryx'. 

INTRODUCTION 
The genus Selaginella P. Beauv. is made up of isophyllous and anisophyllous species. 
The strobili of both the isophyllous and anisophyllous species are terminal, on 
branches and/or branchlets. The anisophyllous species possess one of two basic 
types of strobili depending on whether the distinct anisophylly of the vegetative leaves 
is continued into the reproductive structures or not. 

The tetragonous strobilus has distinct anisophylly of the vegetative leaves which 
is not continued into the reproductive structures; the sporophylls are all uniform or 
subuniform. In contrast, where the anisophylly of the vegetative leaves is continued 
into the reproductive structures, dimorphic sporophylls (Fig. 1b-e) result in a bilateral 
strobilus. 

There are two forms of bilateral strobili — resupinate and non-resupinate strobili. 
In the resupinate strobili (Fig. 1a)the smaller sporophylls are in the same plane as the 
larger lateral vegetative leaves, while the larger sporophylls are in the same plane as 
the smaller median vegetative leaves. The non-resupinate strobili have the smaller 
sporophylls in the same plane as the smaller median vegetative leaves, while the 
larger sporophylls are in the same plane as the larger lateral vegetative leaves. 

This paper reports on the presence of vertical/oblique projections on the adaxial 
surfaces of sporophylls in a number of West African and South American species of 
Selaginella which have been found to possess the bilateral resupinate form of 
strobilus. 

OBSERVATIONS 
During a re-investigation of the West African species of Selaginella it has been found 
that a substantial number of them possess the bilateral resupinate form of strobilus. 
Critical examination of the sporophylls of these species has revealed that the larger 
sporophylls are of an unusual form. They have a vertical/oblique projection on their 
adaxial surfaces. 

Alston (1 959), who gave the first comprehensive account of the genus Selaginella 
in West Africa, listed 20 species. All the 20 species have been examined for the 
presence of this vertical/oblique projection on the adaxial surface of the sporophyll. 
Six of these have tetragonous strobili while 14 have the bilateral resupinate form. The 
projection was found in all the 14species (Table I) possessing the bilateral resupinate 
strobili. 

One hundred and twenty-eight of the 1 33 species of Selaginella listed by Alston et 
al. (1981) in South America have also been examined for this projection. Twenty of 
these have bilateral resupinate form of strobili with the projection on the adaxial 
surface of their sporophylls (Table II). 

•Present address: Botany Department, National Museum of Wales, Cathays Park, Cardiff, 
CF1 3NP. 



50 



FERN GAZETTE: VOLUME 13 PART 1 (1985) 





FIGURE 1. 

a. A bilateral resupinate strobilus of S. molliceps Spring under-surface view (Exell, 500, BM); 

b-c. dimorphic sporophylls of S. leoneensis Hieron. (Harley, F161, BM); b, dorsal sporophyll 

showing complete sporophyll-pteryx and position of ligule in relation to sporophyll-pteryx; c, 

ventral sporophyll; 

d-e. dimorphic sporophylls of S. hartwegiana Spring (from Type specimen, Hartweg 1 477, BM); d, 

dorsal sporophyll showing partial sporophyll-pteryx; e, ventral sporophyll. a, x12, b-e, x30(L = 

lateral leaf, Ig = ligule, M = megasporangia, Ms = megaspore, m = microsporangium, md = median 

leaf, P = sporophyll-pteryx, S = sporophylls, v = midvein). (Arrows indicate the relative positionsof 

the sporophylls to the axis of the strobilus and point to the apex of the strobilus). 



The presence of the projection on the adaxial surface of the larger sporophylls 
gives these sporophylls an unusual and asymmetrical shape. We suggest that the 
projection is the result of a fold of the outer half of the sporophyll onto the adaxial 
surface of itself. It appears that some fusion of the laminal tissue has occurred within 
the fold. About a quarter or so of the folded side of the sporophyll stands out and it is 
this portion that is seen as the vertical/oblique projection on the adaxial surfaceof the 
sporophyll. We propose that this projection be called 'sporophyll-pteryx'; 'pteryx',from 
the Greek, means flap or fold. 

The extent to which the sporophyll appears to have been folded and the extent of 
fusion of its surfaces has resulted in two main forms of sporophyll-pteryx. These we 
are calling partial, and complete, sporophyll-pteryx (Figs. 1d and b). The partial 
sporophyll-pteryx occurs from the apex along about one third the length of the mid- 



SPOROPHYLL-PTERYX IN SELAGINELLA 



51 



vein, then extends obliquely to the edge about midway along the length of the outer 
side of the sporophyll. The fusion of the laminal tissue of the surfaces is incomplete, 
thus the sporophyll-pteryx, in this case, can easily be lifted and moved back to reveal 
parts of the adaxial surfaces of the fold. The complete sporophyll-pteryx occurs from 
the apex along about two-thirds the length of the mid-vein, then extends obliquely to 
the edge of the outer side, as far as the base of the sporophy!'. The fusion of the laminal 
tissue of the surfaces of the folded side is more or less complete except the portion 
forming the sporophyll-pteryx. Any attempt to lift and move the sporophyll-pteryx back 
to reveal parts of the adaxial surfaces of the fold results in damaging the sporophyll. 
Thirteen out of the 14 West African species possess the complete sporophyll- 
pteryx, whilst the remaining one species has the partial sporophyll-pteryx (Table I). 
Seven of the 20 South American species have complete sporophyll-pteryx; the 
remaining 13 species have partial sporophyll-pteryx (Table II). 

TABLE 1. Species of West African Selaginella with sporophyll-pteryx 



S. abyssinica Spring 

S. blepharophylla Alston 

S. buchholzii Hieron. 

S. kalbreyeri Bak. 

S. leoneensis Hieron. 

S. molleri Hieron. 

S. molliceps Spring 



S. protensa Alston 

S. soyauxii Hieron. 

S. squarrosa Bak. 

S. subcordata A.Br, ex Kuhn. 

5. tenerrima A.Br, ex Kuhn. 

S. thomensis Alston 

S. zechii Hieron. 



TABLE II. Species of South American Selaginella with sporophyll-pteryx 

S. moritziana Spring ex Klotzsch P 

S. novae-hollandiae (Sw.) Spring P 

S. pea reel Bak. P 

S. popayanensis Hieron. P 

S. porelloides ( Lam.) Spring C 

S. radiata (Aubl.) Spring P 

S. ramosissima Bak. C 

S. seemannii Bak. P 

S. simplex Bak. C 

S. substipitata Spring P 

C = complete sporophyll-pteryx 
P = partial sporophyll-pteryx 



s 


cavifolia A.Br. 


P 


s. 


cladorrhizans A.Br. 


P 


s. 


flacca Alston 


C 


s. 


flagellata Spring 


C 


s. 


glossophylla Alston ex 
Crabbe & Jermy 


P 


s. 


hartwegiana Spring 


P 


s. 


lychnuchus Spring 


C 


s. 


macilenta Bak. 


C 


s. 


meridensis Alston 


P 


s. 


mollis A.Br. 


P 



DISCUSSION 
Mukhopadhyay and Sen (1981) have reported on the presence of a laminal flap in four 
species of Selaginella - S. bisulcata Spring, S. reticulata (Hook and Grev.) Spring, S. 
tenera (Hook and Grev.) Spring and S. subdiaphana (Wall.) Spring — in India. Reporting 
on the presence of the flap in the Indian species, they stated that "the flap is situated 
next to the ligule" and that "it is a continuation of the lamina and extends from the 
base towards the distal region of the bract orthe sporophyll alongthe midrib". Thisflap 
is the same as the sporophyll-pteryx we are describing in the West African and South 
American species of Selaginella. 



52 FERN GAZETTE: VOLUME 13 PART 1 (1985) 



Our investigation, however, reveals that the sporophyll-pteryx is not situated next 
totheligule(Fig. 1b). Someofthem, upon first glance, appear to be situated next to the 
ligule, for they appear to extend from the apex, along the whole length of the mid-vein, 
to the base of the sporophyll. However, critical examination has shown that they are 
not situated next to the ligule. They do not extend along the whole length of the mid- 
vein but are the same as we have described for the complete sporophyll-pteryx above. 

The possession of sporophyll-pteryx in these species of Selaginella is seen as an 
additional character which can be used in the systematics of the genus Selaginella. 
The presence of the sporophyll-pteryx in 14 West African and 20 South American 
species of Selaginella, in addition to the four species from India, makes it seem highly 
likely that species of Selaginella from other parts of the world will be found to possess 
the sporophyll-pteryx. 

ACKNOWLEDGEMENTS 
The authors are grateful to Mr A.C. Jermy, Miss J. M. Cam us and Miss A.M. Paul of the 
British Museum (Natural History) for the provision of herbarium material and for 
constant encouragement. We are also grateful to Dr D.S. Edwards of the Botany 
Department, University of Cape Coast, Ghana, for sending us collections from Ghana, 
West Africa. 

REFERENCES 
ALSTON, A.H.G. 1 959. The ferns and fern-allies of West Tropical Africa. Suppl. to the 2nd ed. of 

the Flora of West Tropical Africa. VIII + 89 pp. Crown Agents for Overseas Government and 

Administrations, London. 
ALSTON, AH G., JERMY, A.C. & RANKIN, J.M. 1981. The genus Selaginella in Tropical South 

America. Bull. Brit. Mus. (Nat. Hist.) Bot. 9: 223-330. 
MUKHOPADHYAY, R. & SEN, U. 1981. The occurrence of a laminalflap in Selaginella. Fern Gaz. 

12: 180-181. 

BRITISH PTERIDOPHYTE RECORDS 

Compiled by A.J. Worland, BPS Recorder 

The following records have been received and are additions to the Atlas of Ferns 
(1 978); I am grateful to all concerned. As in previous years, the records are presented 
thus; 100km square/ 10km square followed by the recorder's name. WV27 refers to 
Guernsey and WV65 to Jersey (Channel Islands). 

POST 1950 

5.2 Selaginella kraussiana WV27 W. Bennert 

7.4x3 Equisetum * litorale WV27 W. Bennert, 27/60 H. McHaffie & C.N. Page, 37/04 H. 

McHaffie 

10.1 Osmunda regalis WV27 W. Bennert 

13.1 Adiantum-capillus-veneris 13/64 J. Crighton 

1 6. 1 Polypodium vulgare WV65 A.J. Worland 

16 2x1 Polypodium x mantoniae 42/57 R.P.H. Lamb 

16 3 Polypodium australe WV65 A.J. Worland 

1 7. 1 Pteridium aquilinum WV27 W. Bennert, WV65 A.J. Worland 

19.1 Phegopteris connectilis 38/44 D. Welch 

20.1 Oreopteris limbosperma WV65 A.J. Worland 

22. 1 Athyrium filix-femina 48/02 A.O. Chater 
26 2x3 Polystichum * bicknellii WV27 W. Bennert 

26 2 Polystichum aculeatum WV27 W. Bennert 

26.3 Polystichum setiferum WV27 W. Bennert, WV65 A.J. Worland 

27.2 Dryopteris filix-mas WV27 W. Bennert, WV65 A.J. Worland 

27 1x2 Dryopteris * mantoniae 23/60 P.M. Benoit 

27.3 Dryopteris affinis WV65 A.J. Worland 
27.9x8 Dryopteris x deweveri 63/42 A. Willmot 
28. 1 Blechnum spicant 43/33 A. Willmot 



FERNGAZ. 13(1) 1985 53 



ASPLENIUM PUNJABENSE SP. NOV. AND ITS SIGNIFICANCE 
FOR THE STATUS OF CETERACH AND CETERACHOPSIS 



S.S. BIR 
Department of Botany, Punjabi University, Patia!a-147 002, India. 

C.R. FRASER-JENKINS 
c/o Department of Botany, British Museum (Natural History), Cromwell Road, 

London SW7 5BD, England 

and J.D. LOVIS 
Department of Botany, University of Canterbury, Christchurch 1, New Zealand 

ABSTRACT 

A new spleenwort, Asplenium punjabense Bir, Fras.-Jenk. & Lovis and a new 
hybrid, Asplenium * geni-coa/itum Fras.-Jenk. (- A. dalhousiae * A. punjabense), 
are described. A. punjabense is a hexaploid sexual species presumed to be derived 
from A. ceterach subsp. ceterach and A. dalhousiae. Ceterach and Ceterachopsis 
are given subgeneric status and their species are enumerated under the two 
subgenera. Five new specific combinations are made and chromosome numbers 
are recorded for four species of Ceterach for which no previous cytological reports 
exist. The significance of the existence of an allopolyploid species combining 
members of the sub-genera Ceterach and Ceterachopsis is discussed. 

INTRODUCTION 
Ceterach Willd. in the family Aspleniaceae Mett. ex Frank is a small group previously 
treated by most authors as a genus and containing, when circumscribed in a wide 
sense, about twelve morphologically close species (see Copeland 1947 and below). 
From the Himalaya, in addition to Ceterach officinarum DC. (= A ceterach L), the two 
other species usually attributed to Ceterach are C. dalhousiae (Hook.) C.Chr. (= A. 
dalhousiae Hook.) and C. paucivenosum Ching (= A. paucivenosum (Ching) Bir). Of 
these species, the last two, A. dalhousiae and A. paucivenosum, have been further 
separated into another genus or group, Ceterachopsis J. Smith, whose main feature of 
distinction is the presence of an indusium and the absence of scales on the lamina 
unlike in the members of Ceterach sensu stricto. However all three species (as indeed 
can all the species contained within Ceterach and Ceterachopsis) can equally well fall 
within normal definitions of Asplenium and according to Copeland (1 947: 1 69), "It is 
simply impossible to define Asplenium so as to exclude them, and difficult to define 
Ceterach so as to include them". He also stated, "While I entertain no doubt as to the 
propriety of leaving A. dalhousiae in Asplenium its affinity to Ceterach is obvious, and 
makes evident the place in Asplenium {Ceterachopsis J. Smith) from which Ceterach 
was evolved". 

A good deal of discussion has taken place as to the desirability of separating 
Ceterach and other genera from Asplenium, especially in view of the existence of wild 
intergeneric hybrids of Asplenium with Ceterach, Camptosorus and Phyllitis (-Scolo- 
pendrium) (Lovis 1973, Bennert & Meyer 1974). Furthermore, a hybrid between 
Asplenium and Pleurosorus, and another which is in effect trigeneric, combining 
together genomes from Asplenium, Camptosorus and Phyllitis, have both been 
synthesised (Lovis 1973). As a result a number of authors have suggested the merging 
of these and other genera as subgenera within Asplenium; the most important 
discussions are those of Copeland (1 947), for Ceterachopsis, Phyllitis, Asplenidictyum, 
Neottopteris, Ceterach, Camptosorus etc.; Bir (1962, 1963) and Bir in Mehra and Bir 
(1964) for Ceterach and Ceterachopsis; Vida (1963), for Phyllitis and Ceterach; and 



54 FERN GAZETTE: VOLUME 1 3 PART 1 (1985) 



Lovis & Vida (1969), for Phyllitis, Ceterach, Camptosorus and Pleurosorus. Lovis 
(1973), however, puts the case both for and against merging these genera within 
Asplenium, and more recently some important authors working on the Aspleniaceae 
have kept them separate (e.g. Lovis 1 977, Pichi Sermolli 1 977 and Reichstein 1 981 ) 
and it may well be that the presence or absence of intergeneric hybrids in this 
particular family may not be a useful guide to generic limits bearing in mindthe distinct 
morphology of these discrete groups. Clearly though the generic status of all the 
groups is at least open to doubt and they may either be maintained, or reduced to 
subgenera, as has been done for all of them at some stage; Bir(1963)andBir in Mehra 
& Bir (1964: 1 58-1 59), for example, has treated Neottopteris and Asplenidictyum as 
subgenera of Asplenium, even in the absence of intergeneric hybrids, because they 
are considerably closer in morphology to Asplenium than to the other genera 
discussed here and have long been included in it by many authors (see Christensen 
1905-6: 98, 432, Copeland 1947 etc.). More recently, the situation has been 
complicated by the description of Sinephropteris as distinct from Schaffneria (Mickel 
1 976) and by the finding by Kurita (1 972) that Boniniella has n = 76 (x = 38?), in contrast 
to all the rest of the Aspleniaceae which, with the exception of a small groupoftaxa in 
the/4, unilateral complex, uniformly have x = 36. 

In this paper, mainly for the sake of convenience, Ceterach and Ceterachopsis are 
included within Asplenium as subgenera even though the present trend in European 
literature is to maintain Ceterach as a genus. Further discussion follows later in the 
paper as to the alternatives and the separation of the two subgenera from each other. 

OBSERVATIONS AND NEW TAXA 
Bir (1962, 1963; 42-43) and Bir in Mehra and Bir (1964: 158-159) has discussed why 
he followed Christensen (1905-1906) and Copeland (1947) with regard to sinking 
Ceterachopsis (but not Ceterach) within Asplenium. While outlining the evolution of 
various morphological groups within Asplenium, he reported (sub A. ceterach) the 
presence of some Himalayan specimens from Kulu in the Western Himalaya which 
had a morphology intermediate between A. ceterach on the one hand and A. 
dalhousiae on the other. The presence of plants with their morphology thus 
intermediate between Ceterach and Ceterachopsis had potential significance: this 
finding apparently supported Copeland's statements and indicated how Ceterach 
could possibly have evolved from Asplenium through A. dalhousiae. Therefore Bir 
recognised Ceterachopsis as a subgenus of Asplenium, A. dalhousiae Hook, being the 
type species and A. paucivenosum being its only other member then known. More 
recently, however, Love et al. (1977) have followed Ching (1940) in recognising 
Ceterachopsis as a genus. 

Subsequent to Bir's report of intermediates a new spleenwort was discovered by 
J.D. Lovis in a spore sowing made at Leeds from a sheet of Asplenium trichomanes 
subsp. trichomanes collected by Bir from Kulu in the late 1950's. The origin of this 
contaminant, which was clearly distinct from any other Ceterach in culture at Leeds at 
that time was at first mysterious, but investigation subsequently revealed an excellent 
match in a single specimen collected by Bir at the same time in the same locality and 
donated to the BM in the same batch. The source of the stray spores was then obvious. 
This new species showed some characteristics intermediate between/!, ceterach and 
A. dalhousiae and it became clear that it was the origin of Bir's earlier observation (Bir 
1963). Lovis studied its cytology and found the taxon to be hexaploid (n = 108). This 
finding, together with the intermediate morphology, made it probable that it had arisen 
by chromosome doubling in a triploid hybrid of A. ceterach subsp. ceterach (tetraploid, 
n = 72) and Asplenium dalhousiae (diploid, n = 36) both of which are sympatric with it in 
the area. The tetraploid ancestral species, Asplenium ceterach subsp. ceterach, is not 



ASPLENIUM PUNJABENSE & THE STATUS OF CETERACH & CETERACHOPSIS 55 



uncommon, being scattered throughout the slightly drier parts of the Western 
Himalaya, between 1300 and 2700m altitude, extending from Afghanistan to Tehri 
Garhwal, including the Kulu area (Hope 1901-1902). Bir (1959) has also published on 
its presence in the Kulu area. The diploid ancestral species, A. dalhousiae, is 
considerably commoner and is abundant around Kulu, but its distribution extends 
further eastwards into the wetter parts of the Himalaya, from Afghanistan to W & C 
Nepal between 1,000 and 2,500m altitude (see Hope 1901-1902 and also, more 
recently, Mehra & Bir 1957, Bir 1959, 1962, Bir & Shukla 1967 and others). 

Since the new fern was first discovered in a spore culture made from specimens 
collected in the Kulu valley, which was part of the state of Punjab, India at the time of 
collection, it has been given the specific epithet, 'punjabense', but the area nowforms 
part of the state of Himachal Pradesh and is described as such in the citation of 
specimens. It is fortunate that one of us (Fraser-Jenkins) was able to find the species 
growing wild at the type locality as it had previously been known only from the 
spontaneous spore-culture material and single herbarium specimen. Without further 
study in the field, including the making of more wild collections and the discovery of 
other herbarium specimens by Fraser-Jenkins it was not possible to establish that the 
plant was firmly established, rather than of evanescent occurrence, in nature (cf. the 
status of Asplenium eberlei, Eberle 1967, Meyer 1967). 

The diagnosis of the new fern is as follows: 
Asplenium punjabense Bir, Fraser-Jenkins et Lovis, spec. nov. Filix sempervirens; 
morphologia inter Asplenium dalhousiae Hooker et Asplenium ceterach L interjecta; 
differt a specie prima facie abaxiali laminae paleacea et anastomosibus nonnullis 
venarum propter marginem laminae: a specie altera indusiosuppetenteetpaleis facia 
abaxiali laminae minus densis differt; ab utraque specie sporis majoribus atque 
numero chromosomico hexaploido, scilicet paribus chromosomatum meiosi 
regularibus 108 in reproductione sexuali differt. 

An evergreen plant with its morphology intermediate between Asplenium dalhousiae 
Hooker and Asplenium ceterach L It differs from A. dalhousiae by the presence of 
scales on the abaxial (lower) side of the lamina and by several marginal anastomoses 
in its venation; it differs from A. ceterach in the presence of indusia and by the less 
dense scale cover on the abaxial side of the lamina. It differs from both species in its 
larger spores and hexaploid chromosome number with 108 regular pairs of 
chromosomes (bivalents) at meiosis in sexual reproduction. 

Holotype: India (NW Himalaya): Himachal Pradesh, 3km N of Kulu, Beas valley (N of 
Mandi, N of Simla), 1300m alt., among roadside boulders, growing with Asplenium 
dalhousiae, A. trichomanes L. subsp. quadriva/ens D.E. Meyer and Pellaea nitidula 
(Wall, ex Hook.) Bak. Coll.: C.R. Fraser-Jenkins, 6719, 1 /Sept/1 977 (PUN no. 3694). 

Isotypes: ditto (BM, K). 

Paratypes: ditto nos. 6707, 6731 (herb. T. Reichstein, Basel), 6708-671 8, 6720-6730, 
6732-6735 (PUN, PAN, DD, CAL, RAW, KATH, BM, K, PE, CANU, MICH, US); no. 6733 
is also labelled as PUN no. 3693. 

A small, perennial evergreen fern. Rhizome upright or shortly ascendent, densely 
paleate at its apex; scales dark brownish-grey, clathrate, c.4mm long, c. 1 mm wide at 
their base, elongated-triangular in shape with a filiform apex and erose margin; 
mature fronds 3-15 (-20)cm long, c.0.5-2cm wide, forming a compact tuft of fronds; 
stipe dark-brown, thin, usually c.1-2cm long, but longer in plants growing in rock 
crevices, scales at the stipe-base as long as those borne on the rhizome, but becoming 
smaller further up the stipe and on the rachis; lamina up to 15cm long, linear- 



56 



FERN GAZETTE: VOLUME 13 PART 1 (1985) 




FIGURES 1-7. Asplenium punjabense: 1, Habit of plant; 2, Enlargement of pinna lobe showing 
venation and position of sori; 3, Under side of pinna lobe showing the scales; 4, Scales from 
rhizome apex; 5, Scale from stipe; 6, a,b, Scales from under surface of pinna lobe; 7, Spores. 



ASPLENIUM PUNJABENSE & THE STATUS OF CETERACH & CETERACHOPSIS 



57 



lanceolate, pinnatifid or nearly pinnate with deep, ± rounded sinuses between the 
lobes or segments (pinnae), greyish-green or brownish-green, glabrous ontheadaxial 
(upper) side with a few small scales on the rachis, ± densely covered with clathrate, 
brownish, acuminate, erose-margined scales, 0.5-2mm in length on the abaxial 
(lower) side but these do not usually cover the sori completely; the lamina texture is 
herbaceous to subcoriaceous; segments alternate, ovate to oblong, with the margins 
somewhat parallel and obtuse apices; venation forked, subflabellate, with the veins 
partly free and partly anastomosing near the segment margins; sori linear, up to 3mm 
long, slightly hidden by scales, bearing indusia ofthe same shapeand size as the sorus 
and c. 0.3mm wide; spores monolete with the exospore c.42-50jjm long, perispore 
wrinkled, irregularly protruding from the exospore by up to 1 0pm as seen in equatorial 
view. Cytotype: Hexaploid sexual, with 108 regular pairs at meiosis, n= 108. (Figs. 1- 
7). 

Habitat: Apparently without specific soil requirements, on rocks in crevices, or 
between boulders, also along road sides. 

Distribution: Endemic to the Western Himalaya, so far known from the Beas (Kulu) 
valley in Himachal Pradesh, westwards to Kashmir and NW Pakistan (Swat) at 
altitudes from 900 to c. 1,800m. 

Although very closely resembling A. ceterach in habit and frond form, the new 
spleenwort also shows some similarity to A dalhousiae. But in spite of its intermediate 
morphology this fern can be distinguished by the combination of features tabulated 
below: 







Asplenium 


A. punjabense 


A. ceterach 






dalhousiae 




subsp. ceterach 


1. 


Scales on the abaxial 
side of lamina 


absent 


moderately 
dense 


very dense 


2. 


Venation 


free 


partially free 


many veinlet 
fusions 


3. 


Indusium 


present 


present 


absent 


4. 


Spore size 
(exospore length) 


c.28-32gm 


c.42-50um 


c.36-45gm 


5. 


Cytotype 


n = 36 


n= 108 


n = 72 






(diploid) 


(hexaploid) 


(tetraploid) 



Specimens examined 

Asplenium punjabense 

(1) C.R. Fraser-Jenkins 7871-7888 (18 plants). NW Pakistan: Distr. Swat, lower 
Swat valley, c. 5km NE of Mingora, N of Saidu Sharif, 950m alt., 1 /Oct/1 978, stream 
on shale cliff together with Pteris vittata L, Adiantum capillus-veneris L, and 
Cheilanthes pteridioides (Reichb.)C.Chr. (all in BM except 7886 given to T. Reichstein). 

(2) C.R. Fraser-Jenkins 7894-7904, NW Pakistan: Distr. Swat, mid Swat valley, 
c. 4km S of Madyan, N of Saidu Sharif, 1 ,300m. 1 /Oct/1 978, below boulders at road 
side together with Pteris cretica L, Dryopteris nigropa/eacea (Fras.-Jenk.)Fras.-Jenk., 
Asplenium trichomanes L subsp. quadriva/ens D.E. Meyer and Pellaea nitidula (Wall, 
ex Hook.) Bak. (all in BM except 7895 and 7902 given to T. Reichstein, 7901 in PE). 



58 FERN GAZETTE: VOLUME 13 PART 1 (1985) 



(3) S.P. Khullar 244, Kishtwar (Kashmir). Alt. 1,700m. Grows on open dry slopes, 
rare. July/1980 (PUN 3644), with A ceterach and A dalhousiae. 

(4) NW Pakistan: Kulalai Swat, 1,800m. Muquarrab Shah & Dilawar, 377 (ISL); 
Pir Baba, Swat. Muqarrab Shah & Dilawar, 1704 (ISL); Karahar, Swat, 1,290m. 
Muqarrab Shah & Dilawar, 122 (ISL); Kedam, Swat, 1,380m. Muqarrab Shah & 
Dilawar, 630 (ISL); Manglor, Swat, 1,100m. Muqarrab Shah & Dilawar, 543 (ISL); 
Bildgram, Swat, 900m. Muqarrab Shah & Dilawar, 507 (ISL); Sangota, Swat. 
Muqarrab Shah & Dilawar, 1 137 (ISL); Madyan, Swat. A Rahman & /?./?. Stewart, 
15/Aug/1952 (LAH); Dir, 1,410m. A. Rahman, 13/July/1956 (LAH); Swat Ellum, 
Swat, 1,200m. Muqarrah Shah & Dilawar, 28, 18/April/1976, pro parte (ISL). 

A. dalhousiae and A ceterach were also collected by Fraser-Jenkins near to the 
localities for A. punjabense in the Swat valley, and A ceterach also occurs in the Beas 
valley (as well as A. dalhousiae mentioned above). 

One specimen found growing with the population of A. punjabense in the Beas 
valley has abortive spores and its morphology is clearly and recognisably intermediate 
between A punjabense and A dalhousiae. This is therefore described as a new hybrid 
as follows: 

Asplenium x geni-coalitum Fraser-Jenkins, hybr. nov. (= A. dalhousiae * A. 
punjabense). 

Planta morphologia intermedia inter A. dalhousiae et A. punjabense, parum plus 
paleacea ad paginam abaxialam quam in A. dalhousiae. Sporae abortivae. 



- * 



* T », * <■£ 






# r 






B 



m 



.\ 









c 

me 

_. ..._ ^.....w— liy >,. kww»w •iviii v^wi w«, #-«, t-iayim iiui n jju I IJ a UCI I Z> t? , [\UIU VCJIICy, Dll a. II. 

(spontaneous), n = 108. B, A capense, Basutoland, leg. A.F. Braithwaite (AB 28), n = 36 (Phase 
contrast). C, A phillipsianum, Socotra, Gwynne 1 73, n =ca. 72. D, A hauqhtonii, St. Helena, Kerr 

•7Q r, - «^ "70 



FIGURE 9. Diakinesis in spore mother cells, Xc.700. Preparations by J. D.L., all from plants grown 
at the University of Leeds from spores. A, Asplenium punjabense, Kulu valley, Bir s.n. 



79, n = ca. 72. 



ASPLENIUM PUNJABENSE & THE STATUS OF CETERACH & CETERACHOPSIS 59 



Holotype: Same locality, date and collector as the holotype of A. punjabense; no. 6729 
(BM). Cytology unknown. 

This hybrid is named in recognition of its presumably containing genomes from 
Ceterach and Ceterachopsis (as is also true of A. punjabense). 

DISCUSSION 
The taxonomic problems associated with a decision as to the correct systematic 
treatment of Asplenium punjabense are two-fold and individually complex to a degree 
which renders it unsurprising that the three authors of this paper at first found it 
difficult to reach agreement regarding the best solution. 

There is firstly the question of the relationships of Asplenium, Ceterach and 
Ceterachopsis. These three groups may be separated primarily on characters of the 
indusium and frond shape, supported by scale and venation characters. As is 
illustrated in Fig. 8, there are four tenable treatments recognising all three groups as 
genus or subgenus. At one extreme (A), each is recognised as aseparategenus;atthe 
other extreme (D), ultimately chosen here, both Ceterachopsis and Ceterach are 
subordinated to Asplenium as subgenera. The situation of Ceterachopsis is plainly 
intermediate between Asplenium and Ceterach and can with reason, if both these are 
supported as genera, be treated either as a subgenus of Asplenium (C), or as a 
subgenus of Ceterach (D), depending on whether presence/absence of the indusium 
or frond-shape is granted the greater significance. A more drastic treatment (F), 
already in effect adopted by Christensen (1905-06), is to recognise Asplenium and 
Ceterach, but to submerge Ceterachopsis completely within Ceterach, granting it no 
status. 

The second question is that of the proper allocation of A. punjabense. Its 
placement, on morphological criteria, is clearly with Ceterachopsis, so long as this 
group is recognised. However, if our hypothesis regarding the ancestral origin of 
punjabense is correct, then it possesses a majority of genomes from Ceterach, in a 
ratio of 2:1 Ceterach-Ceterachopsis, and is thus logically, in genetic terms, more 
Ceterach than Ceterachopsis!* An instructive parallel here is the relationship of 
Aegilops and Triticum. Contemporary opinion mostly accepts that hexaploid wheat, 
Triticum aestivum, contains one genome (A) from Triticum monococcum and two 
genomes (B and D) from species of Aegilops. Stebbins (1956, p. 238) pointed out that 
on this basis we "... reach the conclusion that the only 'pure' species of Triticum, i.e. 
which does not contain chromosomes derived from Aegilops, is T monococcum ... and 
the maintenance of Triticum and Aegilops as separate genera becomes an absurdity". 
Consequently, most recent authors (e.g. Feldman 1976) submerge Aegilops within 
Triticum. 

Other "intergeneric" allopolyploid species are known in the Aspleniaceae. These 
are Ceterophy/litis hybrida (Milde) Pich.-Serm. (= Phyllitopsis hybrida (Milde) 
Reichstein, Phyllitis hybrida (Milde) C.Chr., Ceterach reichardtii Haracic (Vida 1963, 
Meyer 1981)), Asplenium castaneo-viride Bak. (= A. kobayashii Tag., fide Ching & 
Iwatsuki 1982 (see Lovis, Brownsey, Sleep & Shivas 1973)), Asplenium ebenoides 
Scott and Asplenium pinnatifidum Nutt. (see Lovis 1973, 1977 for review and earlier 
references). For these species independent generic names (Ceterophy/litis Pich.- 
Serm. and Asp/enosorus Wherry) are available. We are unanimous in rejecting such a 
solution for placement of Asplenium punjabense. Such a new compound genus would 
be impossible to characterise satisfactorily. 

*The very close morphological affinity between Asplenium ceterach subsp. bivalens (2x) and A 
ceterach subsp. ceterach (4x) leaves little room for doubt that the tetraploid taxon is either an 
autotetraploid or of segmental alloploid origin from closely related parents — thus all genomes in 
A. ceterach subsp. ceterach must be from Ceterach. 



60 FERN GAZETTE: VOLUME 13 PART 1 (1985) 



There can be little doubt that Ceterach has evolved from a source \n Asplenium by 
changes involving acquisition of scales and a corresponding loss of the (then 
superfluous) indusium, and that, as indicated by Copeland (1 947) and Bir (1 963) the 
source of Ceterach must be sought in Ceterachopsis. Indeed, our observations support 
this concept in as much as the indusium is expressed in A punjabense, despite the 
postulate that it includes four chromosome sets from the ex-indusiate Ceterach. This 
suggests that in Ceterach the indusium is still latent and is indeed suppressed. Of 
course, though punjabense reflects a likely intermediate stage in the evolution of 
Ceterach from the vicinity of Asplenium dalhousiae, it is not itself literally that 
intermediate stage. Its hexaploid chromosome number renders that impossible, since 
it is clear that the evolution of Ceterach from Asplenium took place at the diploid level. 
It might be an autohexaploid derivative of some undiscovered or extinct diploid, but the 
circumstantial evidence, as we have stressed, makes an alloploid origin for 
punjabense much more probable. 

If, as seems almost incontrovertible, Asplenium — > Ceterachopsis — > Ceterach 
constitutes a genuine evolutionary sequence, an interesting conclusion emerges in 
terms of the scales and indusia — to combine Ceterachopsis with Asplenium or with 
Ceterach are both equally natural groupings! It is not unnatural toseparate indifferent 
(but systematically related) taxa two groups that are phylogenetically related, if the 
segregated group constitutes a discrete intact evolutionary unit. The question as to 
whether to recognise and how to relate these groups in a systematic scheme is one 
that has to be resolved on taxonomic and genetical grounds. Fraser-Jenkins would in 
this particular and somewhat special case of Ceterach and Ceterachopsis v. 
Asplenium attach greater significance to the frond shape, in which case Ceterach and 
Ceterachopsis would become more closely grouped together, which is not able to be 
reflected in our choice of ranks when both are subordinated to Asplenium, as here. 

Although the affinity between Ceterach and Ceterachopsis is self-evident, the 
source of Ceterachopsis within Asplenium sensu stricto is not at all clear, and it is 
evident that subordination of Ceterachopsis alone to Asplenium (C & E), though 
taxonomically defensible is, in terms of closeness of relationship, less satisfactory 
than subordination of Ceterachopsis alone to Ceterach (B & F), as advocated by Fraser- 
Jenkins when Ceterach is recognised as a genus. However, in the opinion of all three 
authors, Ceterachopsis is a useful concept and should be retained, although only at 
subgeneric rank within Asplenium, bearing in mind that we also subordinate Ceterach 
to Asplenium. 

There is no doubt that the presence of inter-generic hybrids is an anachronism, 
and ought to be avoided whenever possible. Lovis( 1973) compared the situation in the 
Aspleniaceae with that in the Orchidaceae and Gramineae in as much as in all three 
ill-defined generic limits are a consequence of incomplete and recent activeevolution. 
However, in practical taxonomic terms, these cases are not really similar. To disallow 
intergeneric hybrids in the Orchidaceae and Gramineae would generate drastic 
changes and produce an inconvenient and possibly unworkable taxonomy with 
several vast genera, but the Aspleniaceae is already dominated by the huge genus 
Asplenium, and absorption of all its satellite genera back \n\o Asplenium has relatively 
little effect in terms of practical taxonomy. 

This is a strong argument against accepting Camptosorus, Ceterach, Phyllitis etc. 
as independent genera. However these entities, together with Ceterachopsis, are 
useful names for employment in phylogenetic and biosystematic discussion. We have 
therefore in this paper, for convenience and simplicity, unanimously decided to treat 
both Ceterach and Ceterachopsis as subgenera of Asplenium in order to preserve 
them, a treatment already accepted and preferred by Bir (1 962, 1 963, in Mehra & Bir 



ASPLENIUM PUNJABENSE & THE STATUS OF CETERACH & CETERACHOPSIS 61 



1 964) for Ceterachopsis, and proposed, but not formally adopted, for Ceterach by Vida 
(1 963) and Lovis & Vida (1 969). Fraser-Jenkins strongly advocates though that when 
Ceterach is treated as a genus it is better to treat Ceterachopsis as a subgenus of it. 

SYSTEMATIC SYNOPSIS 
The new grouping of species is as follows: 
Asplenium: 

(a) Subgenus Ceterachopsis (J. Smith ex Ching) Bir in Mehra & Bir, Res. Bull. 
Panjab. Univ., N.S., 15: 159 (1964), 

1. Asplenium dalhousiae Hook., Icon. Plant.: 1. 105 (1837). Type species of the 
subgenus {Ceterach dalhousiae (Hook.) C.Chr., Ind. Fil. 7/ 170 (1905); 
Ceterachopsis dalhousiae (Hook.) Ching, Bull. Fan Mem. Inst. Biol., Bot. 
Ser., 10: 1-22 (1940); Asplenium alternans Wall, ex Hook., Sp. Fil. 3: 92 
(1860)). (Diploid, 2n = 72). 

2. A. paucivenosum (Ching) Bir, Bull. Bot. Surv. India 4: 3 (1962) (Ceterach 
paucivenosa Ching, Bull. Fan Mem. Inst. Biol., 2: 210, t.28 (1931); 
Ceterachopsis paucivenosa (Ching) Ching, Bull. Fan Mem. Inst. Biol., Bot. 
ser., 10: 1 -22 (1 940); A. paucivenosum forma minus Bir, Amer. Fern J. 62: 
46 (1972)). (Tetraploid, 2n = 144. See Bir 1962, 1972). 

3. A. punjabense Bir, Fraser-Jenkins & Lovis. (Hexaploid, n - 108). 

4. A. birii (Love & Love) Bir, Fraser-Jenkins & Lovis, comb. nov. Asplenium 
paucivenosum f. majus Bir, Amer. Fern J. 62: 46 (1972); Ceterachopsis 
birii Love & Love in Taxon 26: 326 (1977). (Octoploid, 2n = 288). The 
relationship of this species to A. magnificum (Ching) Bir, Fraser-Jenkins & 
Lovis requires further study. 

5. A. magnificum (Ching) Bir, Fraser-Jenkins & Lovis, comb. nov. 

(Ceterachopsis magnifica Ching, Bull. Fan Mem. Inst. Biol., Bot. Ser., 11: 
56 (1941)). (Cytology unknown). 

6. A sixth taxon from China has recently been named in the herbarium at 
Peking as Ceterachopsis latiloba Ching & Shing, ined. and appears to be a 
distinct species requiring further study. 

(b) Subgenus Ceterach (Willd.) Vida ex Bir, Fraser-Jenkins & Lovis, stat. nov. 
(Ceterach Willd., Anleit. Selsbstud. Bot.: 578 (1804), nom. cons.). 

1. Asplenium ceterach L. Sp. Plant. 2: 1080 (1753). Type species of the 
subgenus (Ceterach officinarum DC in Lam. & DC, Fl. Franc. 2: 566 
(1805)). 

a. Subsp. ceterach. (Tetraploid, 2n = 144). 

b. Subsp. bivalens (D.E. Meyer) Greuter & Burdet, Willdenowia 10: 17 
(1980). 

(Ceterach officinarum subsp. bivalens D.E. Meyer, Ber. Deutsch. 
Bot. Ges. 77: 8 (1964); Asplenium javorkeanum Vida, Acta Bot. 
Acad. Sci. Hung. 9: 202 (1963)). (Diploid, 2n = 72). 

2. Asplenium aureum Cav., Anal. Cienc. 4: 100 (1801); Descr. 256 (1802) 
(Ceterach aureum (Cav.) L v. Buch., Abh. Akad. Wiss. Berlin 1816-17:361 
(1819). See Benl & Kunkel (1 972)). (Tetraploid, n=72 (Canaries); hexaploid, 
n = 108 (Madeira); octoploid, n = 144 (Canaries)). Evidently a complex 
requiring further taxonomic and cytogenetic study. 

3. Asplenium capense (Kze.) Bir, Fraser-Jenkins & Lovis, comb. nov. 
(Ceterach capensis Kze. Lrnnaea 10: 496 (1836)). (Diploid, n = 36; 
Braithwaite 28, BM. Chase 6822, Umtali, S. Rhodesia (Zimbabwe) is 
tetraploid, n = c.72. Morphologically this collection fits better with this 



62 



FERN GAZETTE: VOLUME 13 PART 1 (1985) 



species than with cordatum SW.). Clearly the capense/cordatum group 
also requires further study. 

4. Asplenium cordatum Sw., Schrad. J. Bot. 1800: 54 (1801) (Acrostichum 
cordatum Thbg, Prod. Fl. Cap.: 171 (1800); Ceterach cordatum (Thbg) 
Desv., Prod.: 223 (1827)). (Tetraploid, n = c.72; Braithwaite 141, BM). 

5. Asplenium haughtonii (Hk.) Bir, Fraser-Jenkins & Lovis, comb. nov. 
(Gymnogramme haughtonii Hk. in Hooker & Baker Synopsis Filicum 381 
(1868)). (Tetraploid, n = 72; St. Helena, Kerr 79). 

6. Asplenium phillipsianum (Kummerle) Bir, Fraser-Jenkins & Lovis comb, 
nov. (Ceterach phillipsianum Kummerle Botan. Kozl. 6: 287(1909); Mag. 
Bot. Lap. 8: 354 (1 909). (Tetraploid, = 72; Socotra, Gwynne 1 73, BM). (This 
species requires further study in order to circumscribe the diagnostic 
features distinguishing it from A. ceterach subsp. ceterach.) 



CETEPACHOPSIS 



CETEEtACH 



L 



ceterachopsi'; 



CETFFtACH 



L 



L 



ASPLFNIUM 



CETERACH 



FIGURE 8. Systematic relationship of Asplenium, Ceterachopsis and Ceterach. Six alternative 
treatments: A, Three separate genera; B & C, Ceterachopsis treated as a subgenus of Ceterach (B) 
ox Asplenium (C); D, Both Ceterachopsis and Ceterach treated as subgenera of Asplenium; E & F, 
Ceterachopsis not recognised, submerged into either Asplenium (E) or Ceterach (F). 



ACKNOWLEDGEMENTS 
We are grateful to Professor R.C. Ching (Peking) for helpful information concerning 
some Chinese species and permission to include details, to Professor Dr T. Reichstein 
(Basel) for fruitful and stimulating discussions, to Dr A.F. Braithwaite (University of 
Nottingham) for providing the collections of Asplenium capense and A cordatum that 
were studied cytologically and to Professor K.U. Kramer (Zurich) for Latin diagnoses. 
We also express our thanks to Dr S.P. Khullar (Chandigarh) for placing at our disposal 
his specimen of A. punjabense collected from Kashmir. Finally, we thank our typist, 
Mrs Anne Robinson of Oxford for the outstanding quality of her work over several 
years. 



ASPLENIUM PUNJABENSE & THE STATUS OF CETERACH & CETERACHOPSIS 63 



REFERENCES 
BENNERT, W. & MEYER, D.E. 1974. Der Gattungsbastard * Asplenoceterach barrancense, hybr. 

nov. (Asplenium majoricum Lit. * Ceterach officinarum Lam. et DC). 
BENL, G. & KUNKEL, G. 1 967. Zur Taxonomie der Gattung Ceterach auf den Kanarischen Inseln; 

Ber. Schweiz. Bot. Ges. 77: 257-265. 
BIR, S.S. 1 959. Cytotaxonomic notes on some Aspleniaceae from Kulu valley; J. Indian Bot. Soc. 

38: 528-539. 
BIR, S.S. 1 962. Taxonomy of the Indian members of family "Aspleniaceae"; Bull. Bot. Surv. India 

4: 1-16. 
BIR, S.S. 1963. Evolution in the Indian Members of genus Asplenium Linn.; J. Indian Bot. Soc. 4: 

41-50. 
BIR. S.S. 1972. Cytological observations on Asplenium paucivenosum; Amer. Fern J. 62: 44-46. 
BIR, S.S. & SHUKLA, P. 1967. Cytology of some North Indian ferns; Cytologia 32: 24-30. 
CHING, R.C. 1 940. The studies of Chinese Ferns, 30; Bull. Fan Mem. Inst. Biol., Bot. Ser., 10:1-22. 
CHING, R.C. & IWATSUKI, K. 1982. Annotations and corrigenda on the Sino-Japanese 

pteridophytes (1). Journal of Japanese Botany 57: 129-132. 
CHRISTENSEN, C. 1905-1906. Index Fi/icum etc. Hafniae. 
COPELAND, E.B. 1947. Genera Fi/icum. Chronica Botanica: Waltham, Mass. 
EBERLE, G. 1 967. Asplenium eberlei D.E. Meyer — eine neue Farnart in der europaischen Flora. 

Natur und Museum 97: 341-346. 
FELDMAN, M. 1976. 'Wheats' in SIMMONDS, N.W. (ed.), Evolution of Crop Plants, pp. 120-128. 

Longman: Harlow. 
HOPE, C.W.W. 1901-1902. The Ferns of North-Western India etc.; J. Bombay Nat. Hist. Soc. 

13(3): 460-461 (1901). 14: 265-266 (1902). 
KURITA, S. 1972. Chromosome numbers of some Japanese ferns (8). Ann. Rep. Foreign 

Students' Coll. Chiba Univ. 7: 47-53. 
LOVE, A., LOVE, D. & PICHI-SERMOLLI, R.E.G. 1977. Cytotaxonomical Atlas of the Pteridophyta. 

Vaduz. 
L0VIS, J.D. 1 973. A biosystematic approach to phylogenetic problems and its application to the 

Aspleniaceae, in A.C. Jermy, J. A. Crabbe & B.A. Thomas (eds.), The phylogeny and 

classification of the ferns; Bot. J. Linn. Soc. 67, suppl. 1 : 21 1-228, et tt. 
L0VIS, J.D. 1977. Evolutionary patterns and processes in ferns, in R.D. Preston & H.W. 

Woolhouse (eds.), Advances in Bot. Research 4: 229-415. London & New York. 
LOVIS, J.D., BROWNSEY, P.J., SLEEP, A. & SHIVAS, M.G. (Mrs Trevor Walker). 1 973. The origin 

of Asplenium balearicum. Brit. Fern Gaz. 10: 263-268 ["1972"]. 
LOVIS, J.D. & VIDA, G. 1 969. The resynthesis and cytogenetic investigation ofxAsp/enophyf/itis 

microdon and * A. jacksonii; Brit. Fern Gaz. 10: 53-67, et tt. 
MEHRA, P.N. &BIR, S.S. 1957. Cytology of some Indian species of genus Asplenium L; Curr. Sci. 

26: 151-152. 
MEHRA, P.N. & BIR, S.S. 1 964. Pteridophytic Flora of Darjeeling and Sikkim Himalayas; Res. Bull. 

Panjab. Univ., n.s., 15: 69-182. 
MEYER, D.E. 1967. Uber neue und seltene Asplenien Europas. Ber. Dtsch-Bot. Ges. 80: 28-39. 
MEYER, D.E. 1981. Phyllitis hybrida (Milde) C. Christensen. Berlin. 1 12 pp. 
MICKEL, J.T. 1 976. Sinephropteris, a new genus of Scolopendrioid ferns. Brittonia 28: 326-328. 
PICHI-SERMOLLI, R.E.G. 1977. Tentamen Pteridophytorum genera in taxonomicum ordinem 

redigendi; Webbia 31: 315-512. 
REICHSTEIN, T. 1981. Hybrids in European Aspleniaceae (Pteridophyta) etc.; Botanica Helvetica 

91: 89- 139. 
STEBBINS, G.L. (1956). Taxonomy and the evolution of genera, with special reference to the 

family Gramineae. Evolution 10: 235-245. 
VIDA, G. 1 963. A new Asplenium (sectio Ceterach) species and the problem of origin of Phyllitis 

hybrida (Milde) Christ; Acta Bot. Acad. Sci. Hung. 9: 197-215. 



64 



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The British Pteridological Society 
THE FERN GAZETTE 

VOLUME 13 PART 1 1985 

CONTENTS 



Page 

MAIN ARTICLES 

A reappraisal of Dryopteris affinis subsp. borreri var. robusta and new records of 
D. affinis subspecies in eastern Europe 

- S. Jessen 1 

Isolating mechanisms in four Himalayan Dryopteris species 

- D. S. Loyal 7 

Anomalous fronds and venation in Polybotrya cervina 

- T. G. Walker 13 

Cytology and taxonomy in Woodsiaceae 

- Y.-L Ma 17 

Ecological observations on the pteridophyte flora of Lantang National Park, 
central Nepal 

- V. L Gurung 25 

Nistarika, a new genus of Polypodiaceae from Silent Valley, south India 

- B. K. Nayar, P. V. Madhusoodanan & M. J. Molly 33 

The generic identity of Po/ypodium banaense 

- W. L A. Hetterscheid 43 

The pteridophyte herbarium of Trinity College Dublin 

- J. Parnell 47 

'Sporophyll-pteryx' in African and American Selaginella 

- N. Quansah &B. A. Thomas 49 

Asplenium punjabense sp. nov. and its significance for the status of 
Ceterach and Ceterachopsis 

- St S. Bir, C. R. Fraser-Jenkins & J. D. Lovis 53 

SHORT NOTE 

Cyrtomium fa/cat urn in Ireland 

- J. P. Cu/linane & C. Crowley 42 

BRITISH PTERIDOPHYTE RECORDS 52 

REVIEW 46 



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FERN GAZ. 13(2) 1986 65 



THE MODULAR GROWTH OF LYCOPODIUM ANNOTINUM 



TERRY V. CALLAGHAN, 

The Institute of Terrestrial Ecology, Merlewood Research Station, 

Grange-over-Sands, Cumbria LA11 6JU, Great Britain 

BRITA M. SVENSSON, 

Department of Plant Ecology, University of Lund, Ostra Vallgatan 14, 

S-223 61 Lund, Sweden 

ALISTAIR HEADLEY 
Department of Botany, University of Manchester, Manchester M13 9PL, 

Great Britain 

ABSTRACT 
Lycopodium annotinum L. is a long-lived plant which consists of a series of annually 
produced segments which can be aged. This paper describes in detail the modular 
construction of the plant and analyses elements of deterministic and opportunistic 
growth by synthesizing a 20-year life cycle from historical records of the plant's 
growth. 

By regarding a plant as a population of modules it ispossibleto usethe methods 
of population biologists for discontinuous variables and the approach of growth 
analysis when studying continuous variables. 

The balance between deterministic and opportunistic growth produces a 
foraging behaviour which enables the plant to harvest patchy resources, and to 
escape interspecific and intraclonal competition. The survival of the plant in its 
unfertile yet relatively predictable environment is enhanced by the recycling of 
nutrients between senescing tissues and growing points. This results in indefinite 
growth and the dominance of vegetative proliferation. Opportunistic colonization of 
new habitats is possible through the subsidized growth of pioneering horizontal 
modules and a constant production of airborne propagules which colonize open 
habitats outside existing locations. "Opportunistic escape" occurs when 
environmental adversity, such as hard substrate or prevented root formation, kills 
dominant growing points resulting in lateral branch proliferation and a great 
extension of the potential zone of exploitation. 

The result is a plant which is successful in both colonizing and surviving in a 
spatially and temporally heterogenous habitat. 

INTRODUCTION 
The sporophyte of Lycopodium annotinum L. (Interrupted Club-moss) is a particularly 
useful tool for investigating population processes and modular growth due to its 
construction of repeated units (segments) within horizontal and vertical branches 
(modules). Seasonality in climate produces markers of annual growth (Callaghan & 
Collins 1 976) which allow a plant to be divided into a series of annually produced units 
which, in cold climates, persist for a long time after death due to slow decomposition 
rates (Jenny et al. 1 949). Roots and strobili can also be regarded as segments and, as 
the plant is long-lived, a time-specific investigation can yield data on growth, 
demography and physiology for a period of over 20 years. 

The advantage of describing a plant as a population of defineable units (Provost 
1978; White 1979, 1984; Harper 1981) is that the methods used by population 
biologists are available. The growth of a plant can then be described in terms of 
numbers of units, the growth of these units and their death, survival and fecundity 
(Kays & Harper 1 974; Callaghan 1 976, 1 984; Bazzaz & Harper 1 977; Maillette 1 982; 
McGraw & Antonovics 1983). 



66 FERN GAZETTE: VOLUME 1 3 PART 2 (1 986) 



It is also possible to interpret interactions between the environment and growth, 
development and physiology of the repeated units in the long term context of the 
population of units within the whole plant (Callaghan 1984; Headley et al. 1985; 
Callaghan et al. 1986). 

Populations of repeated growth units may be studied either by determining age 
class distributions and assuming a stable population to calculate rates of survival 
(Kawano et al. 1 982) or by following a cohort throughout its life cycle (Deevey 1 947; 
Merrell 1947; Harper 1967). The first method fails if the population is unstable 
(Callaghan & Emanuelsson 1985) whereas the second method is impractical with 
long-lived perennials. Lycopodium annotinum presents a plant in which patterns of 
survival can be recorded for a past period by identifying the dates and ages at which 
module death occurs. 

This paper describes in detail the modular growth (or metameric growth, sensu 
White 1984) of this long-lived plant from a stressed environment. The paper serves as 
a basis for detailed investigations of the interactions between the plant and its micro- 
environment (Svensson & Callaghan unpubl.) and long-term age-based physiological 
processes (Headley et al. 1985; Callaghan et al. 1986). 

MATERIALS AND METHODS 
The study was carried out at the Abisko Scientific Research Station on the south shore 
of Lake Tornetrask in Swedish Lapland (68°23'N, 18°55'E). The site was situated in 
the boreal birch forest and was dominated by low density Betula pubescens ssp. 
tortuosa (Ledeb.) Nyman and the ground vegetation consisted mainly of Vaccinium 
vitis-idaea L, V. uliginosum L., Empetrum hermaphroditum Hagerup, Linnaea borealis 
L., and Hylocomium splendens B. & S. Climatic and micro-climatic characteristics of 
the site together with a more detailed description of the vegetation are presented in 
Callaghan et al. (1978) under the heading "Hylocomium site". 

Observations were made in three years: 1975, 1980, and 1982. During 1975, 71 
plants were excavated carefully in the field by tracing the prostrate stems back to a 
point where they were decomposing andtheir integrity had been destroyed. The plants 
were pressed dry and returned to the laboratory for analysis. In 1 980 and 1 982, two 
areas of vegetation 3.8 * 4.2m were mapped in detail, including the exact positions of 
all the horizontal modules of/., annotinum. The plants were then excavated, and after 
measuring the length of each annual increment, and the location of each root, the 
plants were pressed dry for further analysis. 

In the laboratory, each plant was divided into its component segments andthe age 
of each was determined by counting successive morphological markers of annual 
growth (see Callaghan 1 980). The relationship between every annual segment within 
a plant was recorded, as well as dry weight, and number of daughter segments. It was 
also recorded if the segments were dead (i.e. brown with detached microphylls) or 
alive. Of the individual segments analysed in 1975, 1980, and 1982, data from 
approximately 15,000 are presented here. 

Correlations between mean monthly temperatures and annual growth were 
made. To reduce the effect of length variations between segments resulting from 
differences in module size, segment lengths were converted to a length index (I) using 
a method described by Fritts (1976). Variation between segments within modules due 
to positional effects was reduced by omitting the small first segment within each 
module from the analyses, although regressions were not required to remove further 
variation (Fritts 1 976) as recognizable trends of segment length within modules were 
absent. The length index I was calculated as I = Lm/Be where Lm is the measured 
length of a segment and Be is the expected segment length estimated as mean 
segment length per module. 



THE MODULAR GROWTH OF LYCOPODIUM ANNOTINUM 67 



RESULTS 
Organization of the plant and allocation of dry matter 

The sporophyte of L annotinum consists of a series of annually produced segments 
which are aggregated modules growing apically and senescing distally, as the plant 
moves across the ground. Arising from the horizontal segments are smaller, vertically 
orientated photosynthetic segments which are densely covered with microphylls. 
These vertical segments are aggregated into "vertical modules", which are terminated 
either bythedeathoftheyoungest vertical segment inthe moduleor by the initiation of 
a spore-producing strobilus (Fig. 1). Horizontal segments may produce other horizontal 
segments, roots and vertical segments. In contrast, vertical segments can only usually 
produce other vertical segments and strobili. 

The growth of a major plant unit, i.e. a horizontal module (without lateral modules) 
and all of the vertical modules, strobili and roots attached to it, shows a rapid 
accumulation of dry weight reaching a maximum of 3.8g in its seventh year (Fig. 2a). 
Thereafter, weight is lost gradually over a period of more than 17 years. 

The survival of the horizontal module is theoretically indefinite (Fig. 1) but the 
probability of survival of an individual horizontal segment decreases as its age 
increases, which is a familiar biological pattern. 

For the first 2 years, horizontal segments contribute most of the dry weight but 
vertical modules grow rapidly and are the major contributor of biomass for 10 years 
with a peak in year 6 (Fig. 2b). Living horizontal segments survive longer than vertical 
modules and after 13 years horizontal segments are again the major —and ultimately 
the only — contributor of biomass as the vertical modules decompose and become 
detached. The probability of survival of vertical segments varies from that described for 
horizontal segments (Fig. 1) and is described in detail below. Strobili arestrictly annual 
structures and attain dry weights of 18mg (Callaghan 1980). 

Roots contribute a maximum of 5% of the total biomass and reproductive effort in 
terms of dry weight of living strobili reaches only 5. 1 % of the total biomass (Fig. 2b). 

Growth of horizontal modules 

Differentiation of segments. When a new horizontal side-module is formed, the first 
segment to be produced within that module is always smaller in terms of dry weight 
and length than those subsequently produced. For example, the mean length of the 
first segment of modules collected in 1980was 23.4± 1.8(s.e.)mm (n = 223) compared 
with that of other segments which was 68.2 ± 1.7 (s.e.)mm (n = 558). The remaining 
segments show no recognizable differentiation in relation to their order within the 
module. 

Apical dominance. The apex of a main horizontal module of/., annotinum is an active 
and fast growing organ which may photosynthesise (Callaghan 1980). Indeed, it is 
during the first year of growth when roots, vertical modules and other horizontal 
segments are formed (Fig. 1). Growth in this year may reach 170mm and 24mg dry 
weight. This active apex exerts a significant inhibition of the growth of daughter- 
modules such that the effect increases from the oldest to the youngest lateral module 
in a relationship described by:y = 71 - 3.88x (r = 0.64, p< .001, degrees of freedom = 
224) where y = length of one year old segment as percentage of one year old segment 
of dominant branch, andx = order of branch initiation covering 13orders of dominance 
(see Fig. 3 of Callaghan & Emanuelsson 1985). 

At any one time, the population of apices consists of those surviving from the 
previous year, recently dead or dying apices, and newly initiated apices. The 
production of new apical meristems is strongly correlated with the death of dominant 
apices and varies from year to year (Fig. 3). 



68 



FERN GAZETTE: VOLUME 13 PART 2 (1986) 




in 






E 
u 

w 
UJ 

U 

z 
< 

CO 
Q 



Q 

2 => 



O 
O 



FIGURE 1 . Diagrammatic representation of the growth and movement of an unbranched plant of 
L annotinum: For simplicity vertical modules and roots are shown on alternate horizontal 
segments. The lighter hatched areas represent senescing tissues whereas the dotted lines 
contain decomposing tissues also represented by lighter hatching. White breaks represent 
annual markers of growth while the contours denote age classes for the segments. 



THE MODULAR GROWTH OF LYCOPODIUM ANNOTINUM 



69 




SEGMENTS 
live dead 
ROOTS ^^ EM 

STROBILI EB 

VERTICAL 

HORIZONTAL 

OTHERS I I 



SEGMENT AGE (years) 



B w w 




SEGMENT AGE (years) 

FIGURE 2. The dry weight trends (a) and dry weight partitioning (b) in an unbranched plant of L 
annotinum over its life cycle. Bars represent standard errors. The data are based on samples 
collected in 1975 with a sample size of 1079 horizontal segments. 



70 



FERN GAZETTE: VOLUME 13 PART 2 (1986) 



98 



> 


90 


> 




a: 




^ 


R? 


(/) 




LU 




O 


74 


< 




H 




7 




LU 


bb- 


(J 




a: 





Q- 58 



50 




1960 



1965 



1970 
YEAR 



1975 



— i — 
1980 



r24 



■20 



■16 * 

LU 

•12 <* 



8 ° 



FIGURE 3. The relationship between the initiation of daughter-modules (reproduction, thick line) 
and the survival of existing horizontal modules (thin line) over a 22 year period. Data are 
presented for samples collected in 1 975, 1 980, and 1 982. Correlation between annual means: r = 
-0.86 (***with 21 d.f.). 




1980 



FIGURE 4. The relationship between the index of annual horizontal segment length (thin line)and 
the mean monthly temperature (thick line) for the summers of theyears 1955 to 1981. Data are 
based on samples collected in 1975, 1980, and 1982 (n = 1698). See Table 1 for correlation 
analysis. 



THE MODULAR GROWTH OF LYCOPODIUM ANNOTINUM 



71 



Climate. The final length of an horizontal segment is achieved within its year of 
initiation and length is therefore a useful variable to correlate with climate. 

Relationships between mean monthly temperature and segment length show 
significant positive correlations forthe summer months June and July andparticularly 
for the combination of June plus July (Fig. 4). It would appear that 1975 was a 
particularly adverse season both for segment extension growth (Fig. 4) and survival 
(Fig. 3). No correlation was found between survival and precipitation nor between 
survival and temperature. 

Growth of vertical modules 

Differentiation of segments. There are great differences of growth between vertical 
segments according to their roles within the module. The first segment to be produced 
in a vertical module is the smallest whereas the second is the largest (Fig. 5). 
Segments produced after the second show a successive decrease in dry weight (Fig. 5). 
The differentiation mainly results from variations in growth rates during the first 
season of growth. Relative Growth Rates (RGR) of segments 1 to 3 were .021, .047, 
and.049g/g/day respectively (the strobilus had a RGRof .056g/g/day over the same 
period i.e. 17 June to 10 August). 



125 



100 - 



2 75 - 



50 - 



25 



-J 



364 17 



22>1 1-0 



sample size 

144 123 48 64 



frequency 



■8 7-5 



V 

o 

D 
D 


R 
B 



a 



V 

a 
a 

B 


R 
□ 

a 

Q 

a 


1 





2-9 3-9 



V 
B 

a 

a 

n 

D 




R 
B 

a 
a 

B 

n 




1 





25 30 



1-8 



10 



V 

b 

B 

a 

B 
B 

n 


R 
D 

B 
□ 

B 

B 
D 




1 ,J 



0-6 

V 

B 
□ 
B 

B 

n 


0-6 

R 

B 

B 
B 

n 

B 
B 

n 



4 5 6 

BRANCH AGE (years) 



FIGURE 5. Comparison of the dry weights of segments of similar ages and position within 
vegetative (V) and reproductive (R) vertical modu les. Numbers within black bars are the position of 
the segment within the module. The mean dry weight for segment 1 of vegetative and 
reproductive modules was 7 (n = 590) and 1 1.9mg (n = 244) respectively, t = 7.4 (***). The mean 
dry weight of all other segments for vegetative and reproductive modules was 19.8(n = 941) and 
21.4mg (n = 625) respectively (t = 1.86 N.S.). t for differences between segments 1 and the 
remaining segments of vegetative and reproductive modules = 19.5 (***) and 10.8 (***) 
respectively. Data from samples collected in 1975. The frequency of age classes 1 to 7 of vertical 
modules (as percentage of 1644) is also given: the frequencies of age classes younger than 3 
were 48.8% (vegetative) and 0.06% (reproductive). Those for age classes older than 7 years were 
0.7% (vegetative) and 0.42% (reproductive). 



72 



FERN GAZETTE: VOLUME 13 PART 2 (1986) 



TABLE 1. Branching in relation to segment position within reproductive vertical modules, and, 
strobilus weight in relation to age of reproductive modules (standard errors are given in brackets). 



Size of reproductive module (= numbers of vegetative segments plus 
strobilus): 

2 3 4 5 6 7 



>7 



Number of 
modules 
produced per 
vegetative 
segment for 


1 
2 
3 
4 





.294(.117) 
.118(.083) 


.309(.055) 
.423(.057) 
.041(.018) 


.219(.079) 
.828(.116) 
.141 (.05) 



1.3(.13) 
.833(.162) 
.467(.117) 
.67(.047) 


.3(.161) 

1.4(.172) 

7(161) 

.2(.014) 


_ 


segment 
positions: 


5 
6 


— 


— 


— 


— 











strobilus dry 
weight (mg): 


n 


o: 1 


17.7(2.6) 
17 


17.CK.9) 
123 


16.7(1.3) 
64 


14.9(2.0) 
30 


12.6(3.4) 
10 


6.6(2.9) 
5 


Sample size: 



















.4 6.8 



49.2 



25.6 



12 



TABLE 2. Comparison between segments on branches with living apices (n = 263) and dead 
apices (n = 336) of numbers of roots produced (n = 182 and 215, respectively) and the length 
supported by them. Means ± standard errors are presented. 



Length of segment (mm) with: 

roots / segment 

1 root / segment 

2 roots / segment 

3 roots / segment 

Overall length / segment (mm) 
Overall length of segments / root (mm) 

Percentage of segments with: 

roots / segment 

1 root / segment 

2 roots / segment 

3 roots / segment 

Overall number of roots / segment 
Number of roots / dm 2 



BRANCHES WITH APICES 
LIVING DEAD 



48. 8 ± 

71. 2± 

86.6 ± 

131. 0± 


2.6 
3.2 
3.2 
22.7 


50.9 ± 2.2 

71.2 ± 2.4 

101.6±4.9 

1 1 6.8 ± 10.1 


63.5 




65.2 


91.8 




101.9 


46.8 

38.4 

13.7 

1.1 




50.6 

36.0 

12.2 

1.2 



0.69 



0.64 



0.43 



Longest distance between apex and root (mm) 



275 



THE MODULAR GROWTH OF LYCOPODIUM ANNOTINUM 73 



Associated with the differentiation of segments is a pattern of proliferation 
whereby daughter-modules are produced. In all vertical modules, vegetative 
reproductive capacity is minimal in the last segments to be produced independently of 
the ultimate size of the module (Tab. 1). In modules with 3 or more vertical segments, 
branching frequency is greatest in the second and largest module (Tab. 1). However, 
branching is not associated with size as the smallest segments (the first in a module) 
form more new segments than larger segments (i.e. the third, fourth etc.). 
Production ofstrobili. The u Itimate role of a vertical module is to support a strobilus but 
strobili may terminate vertical module growth at any stage. Large modules, i.e. those 
with greater numbers of vertical segments plus a strobilus, are common, and strobilus 
weight decreases as module size increases. Modules composed of 3 vertical segments 
and a strobilus clearly show the greatest frequency and largest strobili (Tab. 1). 
The population of vertical modules. At a given point in time vertical modules attached 
to the same horizontal system may survive and remain vegetative, initiate a strobilus, 
or die. The frequency of strobilus-bearing modules is smaller than that of vegetative 
modules until age class 5 (Fig. 5). Branches consisting of vertical segments with high 
dry weights have a greater probability of producing a strobilus and this is even 
detectable in the small first segments of vegetative and reproductive modules (Fig. 5). 

Vertical modules show an inverted pattern of age-based mortality since the first 
segment to die is the youngest and the last to die is the oldest first segment (Fig. 1 ). This 
is the opposite of that seen in horizontal modules. 

Spore production 

Spore production has been estimated by Plotnikov (1977) as 0.4 x 10 6 per strobilus 
and, with a strobilus density of 4.7 per dm 2 (0.23 ± 0.026 (s.e.)) as recorded in the 
present study, there would be an annual spore production of 1.88 * 10 6 per dm 2 per 
year. The viability of spores is low, only 4% (Svensson unpubl.). 

Root production 

Roots are produced in a very regular way by L an notinum: they are always initiated 
on year horizontal segments and then grow and branch over a four year period. 
Thereafter, deterioration is slow but they may still be functional after 13 years 
(Headley et al. 1985). As the age of the roots increases, the dry weight increases 
relative to the fresh weight, probably due to suberization (Headley et al. 1985). 

At least 1 root is initiated by each horizontal segment but root survival is uncertain 
so that a mean of 0.69 healthy roots per segment was recorded on healthy modules in 
which the apex was active (Tab. 2). Only a small percentage of segments possessed 2 
or 3 healthy roots (Tab. 2). Root weight only reaches 5% of total plant dry weight which 
is at the lower end of the range for evergreens (Shaver & Cutler 1979, Miller et al. 
1982) (Fig. 2b). 

The average length of a horizontal module supported by a root is 91.8mm, and the 
longest distance found between apex and root was 275mm (Tab. 2). 

DISCUSSION 
The genus Lycopodiu m is primitive in terms of life cycle, morphology and anatomy, yet 
highly successful. Lycopodium annotinum and some other members of the genus (L 
se/ago, L. complanatum, L. c/avatum and L alpinum) are successful even in severe 
environments and form an important component of upland boreal and tundra 
vegetation. Their success is based on an ecological strategy enabled by a particular 
balance between opportunistic and deterministic growth (Tomlinson 1982). 

At the first level of organisation, growth of L annotinum is deterministic in that 
the sporophyte is rigidly organized into a predictable pattern of horizontal modules, 
vertical modules, roots and strobili. 



74 FERN GAZETTE: VOLUME 1 3 PART 2 (1 986) 



Horizontal module growth is strongly opportunistic; only the small first segment to 
be produced in a module shows any rigid genetic control although apical dominance 
controls the relative size of segments. Climate has a major effect on the actual size of 
horizontal segments. Survival of the horizontal apex is also opportunistic and is 
controlled by the ability of roots to function. 

Water potential gradients are such that the horizontal apex has the lowest water 
potential and draws water and nutrients from roots as old as 1 2 years which may be 
1 m away (Headley unpubl.). This means that the growth of the apical part of the plant is 
subsidized and does not necessarily reflect its immediate environmental conditions. 
Eventually, however, the subsidized growth of growing points appears to become 
impossible and they die. This then releases other sub-dominant apices from inhibition 
and thereby enables a flexible response by increasing the potential range of habitat 
exploitation (see Callaghan & Emanuelsson 1 985, Fig. 6). This selection of apices by 
the environment helps to control the direction of the plant and when rooting becomes 
possible, a dominant apex extends the clone in a direction dictated mainly by micro- 
topography (Svensson & Callaghan unpubl.). Modules with dead apices had fewer 
roots per cm (Tab. 2) while the horizontal apex has been shown to have depressed 
water potentials on root excision or increased distance from the nearest viable root 
(Headley unpubl.). This lends some support to the hypothesis that a major cause of 
apical death is the inability of roots to grow. 

The proliferation following apical death is most often a response to the micro- 
environment as correlations with climate were not significant. The result is 
opportunistic escape from this unsuitable microenvironment (Tomlinson 1982) and 
may be likened to the foraging behaviour of an animal. It may be argued that the 
"foraging" behaviour of L. annotinum is a successful strategy in a heterogeneous 
environment but that, when favourable microhabitats are found, the plant is at a 
disadvantage since it must grow away from them. However, if L annotinum remained 
in such favourable microhabitats it would soon be over-grown by competitive plants, 
such as the different ericaceous species (Svensson unpubl.). The foraging behaviour is 
therefore successful in a heterogeneous environment and also avoids competition. 
The growth of vertical module segments, in contrast tothat of horizontal modules, 
is deterministic. It is possible to predict which vertical segments will be terminated 
with a strobilus and which will remain vegetative while the production of daughter 
modules from the second vertical segment is regular. However, module size (i.e. the 
number of segments, including strobilus) is not predictable. Vertical modules initiated 
in the same year may carry strobili in different years. The decreasing size of the 
strobilus in relation to increasing module size and height may be related to the 
increasing costs of translocation of nutrients and water (Wall^n 1983), and/orthelack 
of support tissues in this "primitive plant". 

Perennials show a small allocation of dry weight to sexual reproduction compared 
with annuals and biennials. In L annotinum, this allocation reaches a maximum of 
5. 1 % of dry weight which is similar to values quoted by Chester and Shaver (1 982) for 
evergreen and deciduous plants from Alaska and to tundra plants in general 
(Callaghan and Emanuelsson 1985). The probability of successful reproduction is 
reduced even further by a low spore viability of only 4%. However, low spore viability 
and the small allocation of dry matter are compensated for by the vast number of 
spores produced by a strobilus (0.4 x 10 6 ) and the considerable longevity of 
reproducing clones of up to 250years (Oinonen 1968). During 250years, a clone may 
produce 1.5 * 1 0ospores (Callaghan & Emanuelsson 1985). Successful reproduction 
from spores is probably related to the opportunistic colonization of disturbed areas as 
in Viola (Newell 1983). 



THE MODULAR GROWTH OF LYCOPODIUM ANNOTINUM 75 



The death of the apical parts of vertical modules beforethat of the lower segments, 
together with the greater longevity of the horizontal segments, allows the retrans- 
location of elements from senescing vertical segments into the horizontal axis and 
subsequently to new growing points. This transport may be between 63 and 90% 
efficient in terms of N, P and K (Callaghan 1 980). Also, translocation of 1 4C (Callaghan 
1980) was found to be extensive, as in stoloniferous Viola blanda (Newell 1982). Such 
efficient translocation allows the subsidized exploratory growth of the main axis when 
obstacles are encountered. However, should the obstacles be unsurmountable to the 
main axis, there is a flexible and opportunistic response whereby lateral axes are 
initiated and released from suppression with a consequent greatly enhanced zone of 
exploitation. 

In conclusion, the sporophyte of L annotinum shows a balance between 
deterministic and opportunistic growth which 

1) enables the plant to harvest patchy resources from a spatially heterogeneous 
environment via a foraging strategy, 

2) reduces competition within the clone by controlling branching patterns, 

3) reduces competition with other species by producing roots and vertical modules 
in available niches along a constantly advancing axis, 

4) allows survival in impoverished soils by recycling nutrients between segments, 

5) enables varying degress of environmental adversity to be overcome first by 
subsidized growth and then by opportunistic escape following lateral module 
proliferation, 

6) allows survival in a temporally predictable environment by indefinite growth 
associated with vegetative reproduction, and, 

7) enables opportunistic colonization outside existing locations (e.g. following 
forest fires) by constant production of vast numbers of airborne propagules. 

It is not surprising therefore, that this primitive plant is so important ecologically 
throughout the northern latitudes. 

ACKNOWLEDGEMENTS 
The first author is grateful to the Swedish Royal Academy of Sciences for financial 
support of the field work and to Allan Nelson of Merlewood Research Station for 
considerable computer programming assistance. We would also like to thank 
Professor Mats Sonesson, Director of the Abisko Scientific Research Station, for his 
hospitality and the facilities of the research station where the field work was carried 
out. Many of the plant analyses were carried out by staff employed by Cumbria County 
Council's Job Creation Scheme and their help is gratefully acknowledged. Thesecond 
author is grateful to Maria Svensson, Uppsala, Sweden and to Bengt Carlsson, Lund, 
Sweden for their skilful and painstaking fieldwork during 1982, as well as to the 
Swedish Royal Academy of Sciences for financial support. Prof. J.L. Harper, Dr. B. 
Wall^n, and Dr. J. White kindly commented on the manuscript and we are grateful for 
their useful suggestions. 

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Population Biology. Sinauer Association. Mass. USA, pp. 15-47. 



FERNGAZ. 13(2) 1986 77 



THE ECOLOGY OF PTERIDOPHYTES 
IN TASMANIAN COOL TEMPERATE RAINFOREST 



S.J. JARMAN, G. KANTVILAS & M.J. BROWN 

National Parks and Wildlife Service, Magnet Court, Sandy Bay, 

Tasmania 7005, Australia 

ABSTRACT 

Forty nine pteridophytes from 17 families are reported from cool temperate 
rainforest in Tasmania. Epiphytes and terrestrial species are both well represented. 
Their occurrence is influenced by altitude and forest type, but on a regional basis 
most species are widespread. The biogeographical affinities of the rainforest fern 
flora are with New Zealand and southern parts of the eastern Australian mainland. 

INTRODUCTION 
The fern flora of Australia is considered small in relation to the area and latitude of the 
continent (Page & Clifford 1981), containing about 41 6species in 1 18 genera (Jones & 
Clemesha 1981). Only four genera are endemic to Australia (all non-Tasmanian) and 
although six others have a predominantly southern hemisphere distribution most are 
widely dispersed on a global basis. Within Australia, the ferns attain their greatest 
richness in Queensland in tropical rainforests and cloud forests. Generic diversity 
decreases southwards towards Tasmania and also westwards from the coast to the 
drier parts of the continent (Page & Clifford 1981). 

Within Tasmania, ferns andtheir allies arecommon butthey show nopronounced 
specialisation to the Tasmanian environment with only two of approximately 90 
species being endemic. The group is widespread in the state, occurring in most 
vegetation types but reaching its greatest diversity in rainforest, wetsclerophyll forest 
and moist gullies in drier areas. According to Page & Clifford (1981), the number of 
individuals in temperate rainforest and fern gullies in Tasmania and Victoria is very 
striking but their diversity is much lower than could be expected from analogous 
situations in the tropics. Furthermore, they comment that "almost every dominant 
species seems to have a much wider ecological rangethan its tropical counterpart, and 
consequently most occur virtually everywhere that appropriately moist forests and 
fern gullies are present". 

Although the composition of Tasmania's fern flora is well-known (apart from a few 
taxonomic uncertainties), publications which deal directly with the ecology and 
distribution of species are scarce. This publication comprises an account of the ferns in 
Tasmanian cool temperate rainforest and is based mainly on information obtained 
during a broader examination of rainforest in Tasmania (see Jarman et al. 1984). 

METHODS 
The Environment — Cool Temperate Rainforest 

Tasmania is a mountainous island situated approximately 240km south of the south- 
eastern corner of mainland Australia, between latitudes 40° and 43°S. Its rainforest 
vegetation is classified as cool temperate rainforest and forms part of the island's 
Antarctic element (after Hocker 1860) or relict sub-element (after Nelson 1981). 

Rainforest in Tasmania is defined as forests greater than 8m tall, dominated by 
Nothofagus, Eu cry phi a, Phyllocladus, Athrotaxis, Lagarostrobos or Dis el ma (Jarman & 
Brown 1983). It is widespread in the western half of the state with the largest 
unbroken tracts occurring in the northwest. Smaller patches of rainforest occur also in 
the northeast, with isolated pockets scattered elsewhere in the eastern half of the 



78 FERN GAZETTE: VOLUME 1 3 PART 2 (1 986) 



state, usually in gullies. It occurs on both mineral and organic soils, and overlies a wide 
range of rock types from basalts, dolerite and granite to quartzites and sandstones. The 
rainfall requirements of rainforest in Tasmania have been given by Jackson (1 968) as 
1000mm per annum with a summer monthly minimum of 25mm. Rainforest extends 
from sea level to over 1200m in altitude. 

On the basis of floristicandstructural charactertistics, rainforest has been divided 
into four groups (Jarman et al. 1984). Group I (Callidendrous rainforest) is 
characterised by parklike forests with open understoreys, often with a layer of tree 
ferns. The main canopy trees are Nothofagus and Atherosperma, and the diversity of 
woody species in the forest is low. Group II (Thamnic rainforest) also includes forests 
with well-formed trees but they rarely appear parklike because of a shrubby under- 
storey. The diversity of woody species is higher than in Group I and the main canopy 
trees are Nothofagus, Atherosperma, Eucryphia lucida, Phyllocladus, Lagarostrobos 
and/or Athrotaxis selaginoides. Group III (Implicate rainforest) is of lower stature and 
the trees are often of poor form. The understorey is dense with a tangled shrub layer 
from ground level to the canopy. Diversity of woody species is high. The main canopy 
dominants include Phyllocladus, Athrotaxis, Eucryphia and Nothofagus, but 
Leptospermum and Acacia species may also be present. Group IV (Open montane 
forest) is dominated by Athrotaxis cupressoides and comprises low open forests over 
grassy or low shrubby understoreys, the latter usually being less than half the height of 
the forest. The firstthreegroupsformthe bulkof rainforest in Tasmania andthey occur 
from lowland to highland areas. Thamnic and implicate rainforest reach their best 
development in western and southwestern Tasmania whereas callidendrous 
rainforest is best developed in northwestern, central and northeastern Tasmania. 
Open montane rainforest is confined to high altitudes and is found mostly on the 
Central Plateau. 

Sampling 

The survey of pteridophytes was part of a broader survey of rainforest in Tasmania 
(Jarman et al. 1984) involving both phanerogams and cryptogams. Cover and 
abundance data were collected from over 300 quadrats (20 * 20m) which sampledthe 
broad geographic and altitudinal range of rainforest throughout the state. 
Supplementary data were used from Jarman andCrowden(1 978). General comments 
relating to the occurrence of ferns in vegetation other than rainforest are derived from 
literature sources where indicated, from herbarium records or from unpublished 
observations. 

"Mixed forest" (a rainforest understorey below eucalypts —see Gilbert 1 959) was 
not systematically sampled during the survey. However, it is closely related to 
rainforest and is likely to exhibit the same general trends in its fern flora. 

Nomenclature and Taxonomy 

Nomenclature and classification of species follows that given in Jones and Clemesha 
(1981). For convenience throughout the text, the subspecific epithets of Asplenium 
trichomanes subsp. quadrivalens, Cystopteris filix-fragilis subsp. laetivirens and 
Grammitis magellanica subsp. nothofagetii are omitted. 

The taxon referred to in this work as Asplenium cf. terrestre may be either an 
undescribed species or conspecific with A terrestre from New Zealand (P. Brownsey, 
comments with herbarium material, Tasmanian Herbarium). It is widespread in 
Tasmania and has previously been confused with A. flaccidum or A. bulbiferum 
(Brownsey, loc. cit.). 

Grammitis pseudoci/iata has been included in this paper but was not found during 
the survey. This species, previously considered a New Zealand endemic, has only 



ECOLOGY OF TASMANIAN PTERIDOPHYTES 79 



recently been collected in Tasmania (by M. Garrett). Although only found in "mixed 
forest" it has been included because it is epiphytic on Eucryphia lucida, a common 
rainforest dominant. 

RESULTS AND DISCUSSION 
Forty nine species of pteridophytes have been recorded from Tasmanian rainforest 
(see Table 1). These are contained within 15 families of Pteropsida (ferns), one family 
of Lycopsida (clubmosses) and one family of Psilopsida (fork-ferns). Three species, 
Asplenium flabellifolium, Pteridium esculentum and Pteris tremula, appear to be 
adventive and are considered doubtful rainforest species. The remaining 46 species 
represent just over half of the total number of pteridophytes recorded from Tasmania. 
The richest families in rainforest are the Blechnaceae (water ferns) and the Hymeno- 
phyllaceae (filmy ferns). Each is represented by eight species which includes all the 
Tasmanian species of filmy ferns and all but one of the water ferns. Apteropteris 
applanata is the only endemic fern known from Tasmanian rainforest. 

Excluding adventive ferns, the rainforest pteridophytes include 27 terrestrial 
species (including two tree ferns) and 1 9 epiphytes although several species could be 
considered in either group, particularly some of those classified here as epiphytes. 
These two groups can be subdivided using some of the categories from Page's (1 979) 
ecological classification of ferns in mesic and in xeric epiphytic environments in the 
tropics. Thus, three terrestrial habitats can be recognized: the forest floor, streamsides 
and rockfaces. Epiphytic habitats can be divided into two groups: the canopy and the 
understorey. The latter group comprises both crotch and main branch species (after 
Parris 1 976) as well as those occurring below 2m. Low and high climbers, as described 
in Page (1979) for tropical forests, are not a feature of cool temperate rainforest in 
Tasmania with only one species, Microsorium diversifolium, occurring as a climber. 

Terrestrial Ferns 

Diversity among the ground ferns is usually low except in riverine situations, and in 
many forest communities only one or two species are present. Nevertheless, these are 
often abundant and may form a continuous cover up to 1m high. The most common 
ground ferns in undisturbed rainforest are Blechnum watts// and Polystichum 
proliferum. They can occur intermixed but in many forests they are mutually exclusive 
or almost so. They are found throughout Tasmania although the best development of 
Blechnum watts// is on the organic or infertile mineral soils of western and 
southwestern Tasmania whilst Polystichum proliferum is most charactertistic of 
fertile mineral soils in northwestern, central and northeastern Tasmania. Both species 
are found across a range of altitudes, from sea level to treeless alpine situations but in 
extreme highland sites they are confined to sheltered positions among rocks. Thetree 
fern, Dicksonia antarctica, may accompany either species although its greatest 
abundance and most impressive growth is seen in communities where Polystichum is 
present. Dicksonia is particularly common along creek and river margins, and in moist 
gullies or water seepages. However, its occurrence in these situations is probably 
related as much to soil nutrients as to moisture availability since it also occurs in drier 
habitats, particularly on good soils. 

In upland rainforests, particularly in central and northeastern Tasmania, two 
smaller species, Blechnum penna-marina and the facultative epiphyte, Hymeno- 
phyllum peltatum, may be more common on the ground than either B. wattsii or 
Polystichum. Blechnum penna-marina is an obligate soil-dwelling species and occurs 
as small scattered plants, often obscured by litter. Hymenophy Hum peltatum forms low 
dense patches on mossy rocks, logs or exposed roots. A second facultative epiphyte, 
Grammitis billardieri, is often present as small scattered tufts and, like 



80 FERN GAZETTE: VOLUME 13 PART 2 (1986) 



Hymenophyllum peltatum, shows an apparent predilection for mossy surfaces. As 
well as these species, small patches of Lycopodium fastigiatum are sometimes 
present on the forest floor in upland forests. 

Two other ground ferns, Histiopteris incisa and Hypolepis rugosu/a, are 
widespread in rainforest but are most characteristic of disturbed sites. They areableto 
tolerate relatively high light intensities and form a dense cover in open situations 
below breaks in the canopy or along roadsides and tracks. In mature well-developed 
rainforest they occur as occasional plants, often weak and spindly, in local spots which 
are better lit than the surrounding forest. Histiopteris is one of only a few Tasmanian 
ferns which "die down" during the colder winter months. 

Of the remaining forest floor ferns, most are patchy or rare in rainforest. Some of 
the more notable species include Diplazium australe and Pteris comans which are 
locally common on poorly drained soils but are mostly restricted to northwestern 
Tasmania. Gleichenia microphylla also occurs in poorly drained areas, mainly where 
the forest is disturbed, whilst Lindsaeatrichomanoides is sporadic on peaty soil andon 
logs. 

The narrow fringing band of rainforest which occurs along the edges of creeks, 
rivers and lakes represents a separate habitat from the forest floor and often supports 
an abundance of ferns. Many widespread species are present, e.g. Dicksonia 
antarctica, Polystichum proliferum and Blechnum wattsii, as well as several others 
which reach their best development in rainforest in this habitat. For example, 
Blechnum vu/canicum and Sticherus tener are particularly well-developed on the 
steep or overhanging banks of the larger streams although they are not confined to 
these sites. Within rainforest, Blechnum nudum is mostly confined to river edges, 
although outside of rainforest it is widespread in relatively open situations on the 
forest floor. Blechnum f/uviati/e, B. minus and£. chambersii are commonly associated 
with flowing water although occasionally they occur on rockfaces away from water. B. 
chambersii is particularly shade-loving and is rarely present where direct sunlight 
penetrates to the ground surface. 

Rockfaces can also support a wide variety of ferns although at each individual site 
usually only a few species are present. The flora consists mainly of mixtures of 
terrestrial and epiphytic species. However, Cystopteris filix-fragilis, Asplenium 
trichomanes and A. flabellifolium appear to be restricted to this habitat and are 
considered characteristic of rockfaces. The most common terrestrial (including 
streamside) ferns are Blechnum spp. and these are usually represented by scattered 
plants on ledges or anchored in rock crevices. In open, often wet, conditions, 
Blechnum vulcanicum may form locally dense patches with many pendulous wedge- 
shaped fronds. Asplenium bu/biferum and several other less common ground ferns 
are also occasionally found on rockfaces. Epiphytic species present include/4sp/eA?/L/m 
cf. terrestre, Microsorium diversifolium, Tmesipteris billardieri, Lycopodium 
myrtifolium, Grammitis billardieri and Hymenophyllum spp. Hymenophyllum 
flabellatum appears to be very drought tolerant and is often found on the dry roof below 
overhanging rocks. In -high altitude situations in southwestern Tasmania the 
normally host-specific Apteropteris is sometimes abundant on rockfaces, forming a 
dense soft grey mat over the rock surface. 

Epiphytic Ferns 

Epiphytic ferns are widespread in rainforest but are usually smaller and less 
prominent than the ground ferns. The largest of the widespread species are Rumohra 
adiantiformis, Asplenium cf. terrestre and Microsorium diversifolium. Equally 
common but less conspicuous are members of the Hymenophyllaceae and 
Grammitidaceae, e.g. Hymenophyllum rarum, H. peltatum and Grammitis billardieri. 



ECOLOGY OF TASMANIAN PTERIDOPHYTES 81 



Very few of the ferns included in this group are obligate epiphytes and most have been 
found (some rarely) on rocks or soil. Exceptions include Asplenium flaccidum, 
Polyphlebium venosum and Grammitis pseudociliata. 

Host specificity is poor and many vascular species can act as substrates including 
the forest dominants, lesser trees and some of the larger undershrubs. The thick 
fibrous trunks of the tree fern, D/c/rso/7/a, provide excellent substrates and may support 
all of the epiphytic ferns present at any one site. Ferns which appear to be confined to a 
particular host species \r\c\u6e Apteropteris applanata which when epiphytic is found 
only on Athrotaxis trunks (either A. selaginoides or A. cupressoides). Tmesipteris 
elongata and Polyphlebium venosum have been recorded only on Dicksonia trunks in 
rainforest although the latter is also known from the trunks of Cyathea cunninghamii 
in wet sclerophyll forests. Grammitis pseudociliata is known in Tasmania from a single 
collection from Eucryphia lucida. Ctenopteris heterophylla has a wider range but when 
epiphytic shows a preference for Eucryphia lucida or O/earia argophylla. Tmesipteris 
billardieri is found most frequently on tree ferns but also occurs occasionally on the 
mossy buttresses and peaty litter cones of the larger tree species. The conifers, 
particularly Phyllocladus aspleniifolius, are poorly colonized by epiphytic ferns. 

Differences in the fern flora between high and low epiphytic sites are manifest 
mainly in the poor development of the canopy flora. All epiphytes recorded from 
rainforest occur in the understorey but several appear to be very tolerant of a wide 
range of light and moisture conditions and extend into the canopy. However, the 
canopy flora is not characterized by any additional species and thus, in Tasmanian 
rainforest, there are no ferns which can be classed exclusively (or mainly) as high 
epiphytes (cf. Page 1979). Species present in the canopy are typically small ferns 
(Hymenophyllum rarum, H. peltatum, H. cupressiforme, Grammitis magellanica, 
Ctenopteris heterophylla) except for Microsorium diversifo/ium and occasional 
stunted plants of Rumohra adiantiformis. 

Distribution 

Very few ferns are restricted to rainforest in Tasmania. Exceptions include some 
of the uncommon speciessuch as Asplenium flaccidum and Lindsaea trichomanoides. 
Many species are equally abundant in other vegetation types, particularly wet eucalypt 
forests and some, including Asplenium bulbiferum, Lastreopsis acuminata, 
Gleichenia microphylla, Todea barbara and Cyathea australis (possibly also Sticherus 
lobatus and Blechnum patersonii) are developed better outside of rainforest than 
within it. 

Within rainforest, the distribution of ferns varies among the main rainforest 
groups. Callidendrous rainforest possesses the richest and most luxuriant fern flora 
(although this is not necessarily true at every site). It contains most of the common 
terrestrial and epiphytic ferns, but is especially typified by Polystichum and Dicksonia 
which dominate the understorey vegetation in many medium to low altitude forests. 
Two species are restricted to callidendrous forests, viz. the tree fern, Cyathea australis 
which, in rainforest, is confined to small relict patches mostly on the east coast, and 
Tmesipteris elongata. Several other species such as Hymenophyllum cupressiforme 
and species mainly epiphytic on tree ferns, e.g. Hymenophyllum flabellatum, 
Polyphlebium venosum and Tmesipteris billardieri, are found most commonly in 
callidendrous forest although they occur also in other rainforest groups. 

Most fern species which are found in callidendrous forests occur also in thamnic 
rainforest but they rarely attain the same luxuriance. Blechnum wattsii, the main 
ground fern, is an exception and its leathery pinnate fronds may form a dense dark 
green cover up to 1m high, particularly wherethe canopy is broken. The most common 
epiphytes are Hymenophyllum rarum and Grammitis billardieri. The decline in 



82 FERN GAZETTE: VOLUME 1 3 PART 2 (1 986) 



luxuriance of the fern flora compared with that in callidendrous forests may result, in 
part, from the scarcity of the "popular" epiphytic substrate, Dicksonia. However, it may 
also reflect some more fundamental habitat difference, possibly one associated with 
humidity. 

In general, ferns are poorly developed in implicate rainforest although Blechnum 
watts// is common in several communities. Many characteristic species from 
callidendrous and/or thamnic forests are absent or rare including Dicksonia (and its 
epiphytes), Rumohra adiantiformis, Asplenium cf. terrestre, Microsorium 
diversifolium and Ctenopteris heterophylla. However, Grammitis billardieri, Hymeno- 
phyllum rarum and H. peltatum are widespread and H. australe may be present in 
moister situations. Hymenophyllum marginatum reaches its best development in 
implicate rainforests although it is easily overlooked because of its small size and 
resemblance to some liverworts. It is absent from callidendrous forests and most 
thamnic forests (except riverine situations) but sometimes occurs in open montane 
forests. 

Open montane rainforest contains an impoverished fern flora. Grammitis 
poeppigiana is restricted to this group and occurs in small patches usually in rock 
crevices. Apteropteris applanata (on Athrotaxis trunks) and Hymenophyllum peltatum 
(epiphytic or on mossy rockfaces) may be locally common. Other ferns are rarely 
present and are confined to the most sheltered, shaded microhabitats. 

Although most ferns are widely dispersed throughout the state, several show 
restricted distributions. Lindsaea trichomanoides has been recorded only from south- 
western Tasmania and Grammitis pseudociliata is even rarer, being known from only 
one site in southern Tasmania. Lastreopsis hispida, Tmesipteris elongata, Pteris 
comans, Diplazium australe and Sticherus lobatus are local and are found mainly in 
northwestern Tasmania although rare occurrences are known elsewhere. Blechnum 
patersonii is found mostly in moist gullies in northern or eastern parts of the state and 
Cystopteris fi/ix-fragi/is occurs on shaded rockfaces in central and southwestern 
Tasmania. A number of other species show wider distributions but are constrained by 
specialised habitats. For example, the distribution of Asplenium trichomanes is 
contained within areas where limestone outcrops and the distribution of Apteropteris 
applanata (in its epiphytic form) is restricted to that of its host, Athrotaxis. Some 
species, although more widely dispersed in the state, are uncommon, e.g. Asplenium 
flaccidum and Lycopodium myrtifolium. 

As a general trend, the abundance and luxuriance of ferns declines with 
increasing altitude with the most pronounced effect being apparent at 600- 700m 
above sea level. Several of the larger ground ferns, e.g. Polystichum proliferum and 
Blechnum wattsii, may be present in rainforest above about 700m but they are 
represented by scattered, often small individuals which occur mostly in sheltered 
sites. A few species, including Hymenophyllum peltatum, Grammitis billardieri, 
Blechnum penna-marina and Lycopodium fastigiatum, are well-adapted to high 
altitude situations although not necessarily restricted to them. In montane rainforests 
(above 1000m), Grammitis billardieri is replaced by G. poeppigiana which occurs in 
small dense mats in rock crevices. Cystopteris fi/ix-fragi/is also appears to be restricted 
to upland forests (altitudes above 800m). 

A similar trend of altitudinal zonation is apparent in Victorian cool temperate 
rainforest (see Howard & Ashton 1973) although the altitude differences occur at 
lower elevations in Tasmania. One species, Hymenophyllum peltatum, which was 
found only in high altitude forests in Victoria (Howard & Ashton 1 973) occurs across a 
wide range of altitudes in Tasmanian rainforest and may be present at sea level in 
some riverine communities in western parts of the state. 



ECOLOGY OF TASMANIAN PTERIDOPHYTES 83 



Biogeography 

The geographical distribution of ferns found in Tasmanian rainforest, taken from 
Jones and Clemesha (1981), is shown in Table 1. Over two thirds of the rainforest 
ferns are confined to Tasmania-Australia, Tasmania-New Zealand or Tasmania- 
Australia-New Zealand. Several additional species occurring in these regions also 
extend their range into the Pacific Islands. Other biogeographical elements are poorly 
represented but include a few Cosmopolitan, Endemic, Old World Tropics and Austral 
species (after Parris 1 976). A comparison between Tasmania and other land masses 
containing cool temperate rainforest indicates that 8% of Tasmanian rainforest ferns 
are shared with South America, 69% are shared with New Zealand and 92% are 
shared with mainland Australia. 

The southern affinities of the Tasmanian rainforest ferns are borne out by their 
occurrence within Australia. Only 1 1 of the 49 species (22%) are found in tropical 
northern Queensland whereas 57% are found in southern Queensland. Eighty six per 
cent occur also in New South Wales and in Victoria, the most southerly part of the 
Australian mainland. These figures are reduced, at least with respect to Victoria, 
where the comparison is restricted to the same habitat in both states. Thus, of the 40 
species listed by Howard and Ashton (1973) in cool temperate rainforest in Victoria, 31 
were recorded in Tasmania during the present survey. Thus, 53% of species from 
rainforest habitats are shared between Tasmania and Victoria. The most notable 
Tasmanian species absent from Victorian rainforest include Asplenium cf. terrestre, 
Sticherus tener, Hymenophyllum marginatum and Grammitis magellanica. 

SUMMARY 
The fern flora in Tasmania's rainforest is small and comprises 40 species (including 
three species which may be adventive). Afewferns are confinedto rainforest but most 
occur also in wet eucalypt forests or scrub. The most common ferns include 
Polystichum proliferum, Blechnum watts//, Dicksonia antarctica, Rumohra 
adiantiformis, Microsorium diversifolium, Asplenium cf. terrestre, Hymenophyllum 
rarum and Grammitis billardieri but many other species are widespread or locally 
common in particular situations. Some of the rarer ferns include Lindsaea 
trichomanoides, Cystopteris filix-fragilis, Grammitis pseudociliata and Lastreopsis 
hispida. Both epiphytic and ground species (including two tree ferns) are represented. 
In most communities, ground ferns account for the greatest biomass within the fern 
flora whilst epiphytes account for the greatest diversity. Only one endemic fern occurs 
in Tasmanian rainforest. 

As a general trend, ferns decline in prominence and abundance from lowland 
situations to high altitudes. They also show differences in luxuriance and diversity 
among the four rainforest groups, with their best development in callidendrous 
rainforest. They are most impoverished in high altitude open montane rainforests. 

Over two thirds of the ferns found in rainforest are confined to Australasia 
(Tasmania, Australia and New Zealand). Within Australia, the rainforest fern floras of 
Tasmania and Victoria are very similar with over thirty species being common to the 
rainforests of both states. 



84 



FERN GAZETTE: VOLUME 13 PART 2 (1986) 



TABLE 1. Habitat and distribution of ferns recorded from Tasmanian rainforest. Distributions 
outside of Tasmania are taken from Jones and Clemesha (1 981). (Abbreviations: N = New South 
Wales, NT = Northern Territory, NZ = New Zealand, Q = Queensland, SA = South Australia, SAf = 
SouthAfrica, SAm = South America, Tas = Tasmania, V = Victoria). 



Species 



Habitat and Distribution 



Apteropteris applanata On rocks or epiphytic (on Athrotaxis), scattered throughout 

A.M. Gray & R.G. Williams western, central and southwestern Tasmania; endemic. 

Mostly on soil or rocks but uncommon in rainforest; more 
widespread in wet sclerophyll forest, particularly in northwestern, 
northern, and eastern parts of the state. Q, N, V, SA, NZ, Asia. 

Uncommon in rainforest and probably adventive; mostly found 
among rocks in drier forests. All states, NZ. 

Epiphytic, scattered in rainforest throughout Tas. Q, N, V, NZ, 
Pacific Is. 

Restricted to limestone rocks; present in several vegetation 
formations. All states except Q, NZ, Europe. 



Asplenium bulbil erum 
Forst.f. 



A. flabellifolium 
Cav. 



A. flaccidum 
Forst.f. 



A. trichomanes L. 

subsp. quadrivalens 
D.E. Meyer 

A. cf. terrestre 

Blechnum chambers// 
Tindale 

B. fluv/at/fe (R.Br.) 
E.J. Lowe ex Salom. 

B. minus (R.Br.) Ettingsh. 

B. nudum (Labill.) Mett. 
ex Luerss. 

B. patersonii (R.Br.) Mett. 

B. penna-marina (Poir.) 
Kuhn 

B. vu/can/cum (Bl.) Kuhn 

B. watts// Tindale 

Ctenopter/s heterophylla 
(Labill.) Tindale 

Cyathea austral is (R.Br.) 
Domin 



A widespread epiphyte in rainforest. 

Ground species of shady situations in wet forests mostly along 
rivers or drainage channels but also on rockfaces. 
Q, N, SA, NZ, Fiji. 

Widespread ground species in wet forests near creeks but also on 
rockfaces. N, V, NZ. 

Common ground species along creeks. Q, N, V, SA, NZ. 

Widespread ground species; in rainforest, mostly along edges of 
creeks and rivers where the canopy is broken. Q, N, V, SA. 

Ground species; local, near creeks in wet forests in northern and 
eastern Tasmania. Q, N, V, Fiji. 

Widespread ground species in high altitude situations in several 
vegetation formations; occasionally found to sea level. 
N, V, NZ, SAm, SubAntarctic Is. 

Ground species, most common along the edges of creeks and rivers 
in western and southwestern Tasmania; also on rockfaces. NZ. 

Widespread ground species in wet forests. Q, N, V, SA. 

Epiphytic or on rocks; scattered in wet forests but also extending 
into drier vegetation. V, NZ. 

Tree fern; occasional in rainforest but common in wet eucalypt 
gullies in eastern, northeastern and northern Tasmania. Q, N, V. 

Occasional on rocks in central and southwestern Tasmania in high 
V, NZ. 



Cyst opt er is fi/ix-fragi/is 

(L.) Bernh. subsp. laetivirens altitude forests. N 

(Prent.) C.Chr. 

Dicksonia antarctica Tree fern; widespread in wet forests. Q, N, V, SA 

Labill. 

Dip/azium australe 
(R.Br.) N.A. Wakefield 

Gleichenia microphylla R.Br. Ground fern; occasional in rainforest, widespread in damp 
situations in other wet forests. All states, NZ. 

Epiphytic or terrestrial; widespread in wet forests. 
N, V, NZ, SAf, SAm. 

Scattered in wet forests, mainly epiphytic. NZ. 



Ground fern; patchy in wet forests, mainly in northwestern 
Tasmania. Q, N, V, NZ. 



Grammitis billardieri Willd. 



G. magellanica Desv. 

subsp. nothofagetii 
Parris 



ECOLOGY OF TASMANIAN PTERIDOPHYTES 



85 



G. poeppigiana 
(Mett.) Pic. Ser. 

G. pseudociliata 
Parris 

Histiopteris incisa 
(Thunb.) J.Sm. 

Hymenophyllum australe 

Willd. 

H. cupressiforme Labill. 

H. flabellatum Labill. 

H. marginatum Hook. & 

Grev. 

H. peltatum (Poir.) Desv. 



H. rarum R.Br. 

Hypolepis rugosula 
(Labill.) J.Sm. 

Lastreopsis acuminata 
(Houlston) Morton 

L hispida (Sw.) Tindale 

Lindsaea trichomanoides 
Dryand. 

Lycopodium fastigiatum 
R.Br. 

L myrtifo/ium Forst.f. 

Microsorium divers ifo Hum 
(Willd.) Copel. 

Polyphlebium venosum 
(R.Br.) Copel. 

Polysticnum proliferum 
(R.Br.) Presl ' 

Pteridium esculentum 
(Forst.f.) Cockayne 

Pteris comans 
Forst.f. 

P. tremula 
R.Br. 

Rumohra adiantiformis 
(Forst.f.) Ching 

Sticherus lob at us 
N.A. Wakefield 

S. tener (R.Br.) Ching 

Tmesipteris billardieri 
Endl. 



T. el on gat a Dang. 

Todea bar bar a 
(L.) T. Moore 



Scattered in high altitude vegetation, usually in sheltered rock 
crevices. N, V, NZ, SubAntarct. Is. 

Epiphyte known from one location only in Tasmania. NZ. 



Ground fern; widespread in wet forests. Q, N, V, SA, NT, Tropics & 
S. Hemisphere. 

Common in wet forests, mostly on buttresses or mossy rocks 
where the humidity is high. Q, N, V. 

Usually epiphytic; widespread in wet forests. Q, N, V. 

Mostly an epiphyte of Dicksonia, but also on logs or rocks. 
Q, N, V, NZ, Pacific Is. 

Mostly epiphytic in western and southwestern Tasmania, in wet 
scrub and forests. Q, N. 

Widespread epiphytic or rock species in wet forests; common at 

high altitudes but also extending to sea level. 

Q, N, V, NZ, SAf, Mascarene Is. (Also in S. Chile — Looser 1 948). 

Widespread in wet forests; mainly epiphytic. N, V, NZ. 

Widespread ground fern in wet forests. Q, N, V, SA, NZ. 

Ground fern; uncommon in rainforest but more widespread in 
other wet forests. Q, N, V, SA. 

Ground fern; sporadic, mostly in northwestern Tasmania along 
creeks. N, V, NZ. 

Localized in rainforests in southern and southwestern Tasmania, 
mostly along the larger rivers. N, V, NZ. 

Widespread ground species in upland vegetation. Q, N, V, NZ. 

Occasional in wet forests; epiphytic or terrestrial. Q, N, V, NZ. 

Widespread across a range of vegetation types including dry 
forests; epiphytic or terrestrial. Q, N, V, NZ, Norfolk Is. 

Epiphyte; restricted to Dicksonia antarctica trunks in rainforest but 
also on Cyathea cunninghamii trunks in wet eucalypt forests. 
Q, N, V, NZ. 

Widespread ground fern in wet forests. N, V, SA. 

Ground fern; adventive in rainforest, common in other wet and dry 
lowland vegetation. All states, NZ, Pacific Is. 

Ground fern; patchy in rainforest, mainly found in northwestern 
Tasmania. Q, N, V, NZ, Pacific Is. 

Uncommon ground fern in Tasmania, probably adventive in 
rainforest. Q, N, V, SA, NT, NZ, Norfolk Is, Fiji. 

Widespread terrestrial and epiphytic species. Q, N, V, NZ, SAm, SAf 

Ground fern; patchy in rainforest, mostly in forests from north- 
western Tasmania. Q, N, V. 

Widespread ground fern in wet forests, often found along rivers or 
road cuttings. N, V. 

Widespread in wet forests, usually epiphytic on Dicksonia 
antarctica but occasionally in peat or on logs. N, V. 

Recorded only from Dicksonia trunks; occasional in lowland 
Tasmania but possibly overlooked. V, NZ. 

Ground fern; uncommon in rainforest, found elsewhere along 
creeks and in gullies mostly in drier lowland parts of the state. 
Q, N, V, SA, NZ, SAf. 



86 FERN GAZETTE: VOLUME 1 3 PART 2 (1 986) 



ACKNOWLEDGEMENTS 
We wish to thank M. Garrett and M. Neyland for their comments on the manuscript 
and M. Garrett for permission to cite his unpublished record of Grammitis 
pseudociliata. The work was undertaken with a grant provided by the Forest Ecology 
Research Fund. 

REFERENCES 
GILBERT, J.M. 1 959. Forest succession in the Florentine Valley, Tasmania. Pap. Proc. Roy. Soc. 

Tas. 93, 129-151. 
HOOKER, J.D. 1860. Introductory Essay. In Botany of the Antarctic Voyage of H.M. Discovery 

Ships "Erebus" and "Terror" in the years 1839-1843, Vol. 3, Flora Tasmaniae, Reeve, 

London. 
HOWARD, T.M. &ASHTON, D.H. 1973. The distribution of Nothofagus cunninghamii rainforest. 

Proc. Roy. Soc. Vic. 86: 47-76. 
JACKSON, W.D. 1968. Fire, Air, Water and Earth — an Elemental Ecology of Tasmania. Proc. 

Ecol. Soc. Aust. 3: 9-16. 
JARMAN, S.J. & BROWN, M.J. 1983. A definition of cool temperate rainforest in Tasmania. 

Search 14(3-4): 81-87. 
JARMAN, S.J., BROWN, M.J. & KANTVILAS, G. 1 984. Rainforest in Tasmania. National Parks 

and Wildlife Service, Tasmania. 
JARMAN, S.J. &CROWDEN, R.K. 1 978. A Survey of Vegetation. Lower Gordon Scientific Survey. 

Hydro Electric Commission, Tasmania. 
JONES, D.L &CLEMESHA, S.C. 1981. Australian Ferns and Fern Allies. 2nd Edn. A.H. &A.W. 

Reed, Sydney. 
LOOSER, G. 1948. The ferns of southern Chile. American Fern Journal 38: 71-87. 
NELSON, E.C. 1981. Phytogeography of southern Australia. In A. Keast (ed.), Ecological 

biogeography of Australia. W. Junk, The Hague. 
PAGE, C.N. 1979. The diversity of ferns. An ecological perspective. In A.F. Dyer (Ed.), The 

Experimental Biology of Ferns, Academic Press, London. 
PAGE, C.N. & CLIFFORD, H.T. 1 981 . Ecological Biogeography of Australian Conifers and Ferns. In 

A. Keast (Ed.), Ecological Biogeography of Australia. W. Junk, The Hague. 
PARRIS, B.S. 1976. Ecology and biogeography of New Zealand pteridophytes. Fern Gaz. 1 1(4): 

231-245. 



FERN GAZ. 13(2) 1986 87 

TMESIPTERIS IN VANUATU (NEW HEBRIDES) 



A.F. BRAITHWAITF 
Botany Department, University Park, Nottingham NG7 2RD 

ABSTRACT 

Cytological, anatomical and ecological observations on Tmesipteris from Vanuatu 
are presented and attention is drawn to the variation in stem anatomy and ecology 
in the genus as a whole. Three species are recognised in the archipelago: T. oblongi- 
folia sp. nov. (n = 104) from Tanna and Aneityum, T. vanuatensis sp. nov. (n = 104) 
from Espiritu Santo and T. oblanceolata (R.Br.) Desv. (n = 208) from Aneityum and 
Espiritu Santo. 

INTRODUCTION 
The genus Tmesipteris was based by Bernhardi (1801) on the species, Lycopodium 
tannense, which was described a year earlier by Sprengel (1800) from a Forster 
collection supposedly made on Tanna, Vanuatu. However, no Forster specimen from 
Tanna has ever been found and Sprengel's description is very brief and only sufficient 
to placethe species in the genus. Tmesipteris was collected again in the archipelago by 
both Milne and Seeman in the middle of the last century but these collections came 
from Aneityum and differed from Bernhardi's description and illustration. 
Consequently the identity of the type species, T tannensis (Spreng.)Bernh., remained 
for many years in doubt and our knowledge of the genus in the islands as a whole has 
been based on very few specimens and was inevitably incomplete. 

Recently Tmesipteris has been found again on Tanna, Aneityum and, for the first 
time, on Espiritu Santo. The new material from Tanna has already clarified the 
typification and identity of T tannensis. It differs from the species illustrated by 
Bernhardi (1801) and Chinnock (1 976) has provided convincing evidence to show that 
the Forster specimen described and illustrated by Bernhardi in fact came from Dusky 
Sound, New Zealand, and has suggested that the reference to Tanna by Sprengel was 
probably an error. The Bernhardi illustration is, therefore, selected as the neotype of L 
tannense Spreng. and the type species is now considered to be restricted to New 
Zealand. 

The present paper gives a more complete account, including cytological and 
anatomical studies, of Tmesipteris in Vanuatu. The material from Tanna belongs to the 
T. lanceolata group and is described as a new species, T oblongifolia, which is also 
found on Aneityum. Two further species are also recognised in the archipelago, 
namely, T vanuatensis A.Braith. a new species proposed for material from Espiritu 
Santo, and T oblanceolata (R.Br.) Desv. from Aneityum and Espiritu Santo. 

MATERIALS AND METHODS 
The material was collected by the author during the 1971 Royal Society and Percy 
Sladen Expedition to the New Hebrides. The details of the localities of the collections 
are given in Table 1. At each locality plants were collected for herbarium specimens 
and preserved in 70% alcohol for anatomical studies. When available, sporangial 
material was also fixed in the field in 1 :3 acetic-alcohol and despatched by air to the 
U.K. where it was stored in a deep freeze. Meiotic preparations for chromosome counts 
were subsequently made using the acetocarmine squash method. The material for 
anatomical studies was embedded in paraffin wax and the sections stained in saf rani n 
and light green or aniline blue. Spore samples were taken from dried herbarium 
specimens and mounted in gum chloral for measurement. Herbarium material of each 
species is deposited in the Herbarium, Royal Botanic Gardens, Kew. 



88 FERN GAZETTE: VOLUME 1 3 PART 2 (1 986) 



CYTOLOGY 
The results of the chromosome counts are summarised in Table 1 and representative 
cells are illustrated in Figs. 1 & 2. It is difficult to produce absolutely unequivocal 
counts in the genus because of a) the high chromosome numbers, b) variation in the 
size of chromosomes and c) the sometimes peculiar shapes of the bivalents. The latter 
has generally been attributed to a laxity in the spiral structure (Manton 1950, Lovis 
1977). Nevertheless the counts listed in Table 1 are accurate to within one or two 
chromosomes except where indicated, and even in these cases the order of the 
chromosome number is not in doubt. 

TABLE 1. Chromosome Numbers in Tmesipteris from Vanuatu 

Species Locality Chromosome Spore length* Length of 

number (urn) Stomata* (jum) 

T. oblong/folia RSNH 2146, Woptiabo, 

Aneityum 

RSNH 2204, Mt. Toukosmeru, 
Tanna 

RSNH 2211, Mt. Toukosmeru, 
Tanna 

T. vanuatensis RSNH 2354, Apouna Valley, 

Espiritu Santo 

RSNH 2382, Mt. Tabwemasana, 
Espiritu Santo 

T. oblanceolata RSNH 2112, Inrero, 

Aneityum 

RSNH 2152, Nezwon Nelgon, 
Aneityum 

* Data based on 100 measurements from one specimen from each collection. 

The numbers recorded here fall intothecytological pattern already established for 
the genus by Barber (1 957) from Australian material of two groups with n = 104and n = 
208 respectively. These relatively high numbers have in the past been designated as 
various levels of polyploidy but are here interpreted in terms of the lowest extant 
number known in the Psilotales, which is n = 52 (Lovis 1 977). Thus T. oblongifolia and 
T. vanuatensis are tetraploids and T. oblanceolata is an octoploid. 

The mean length of spores and stomata are also given in Table 1 . It is clear from 
this data that the spores and stomata of the octoploid, T. oblanceolata, are considerably 
larger than those of the tetraploids, T. oblongifolia and T. vanuatensis. Although the 
samples are small, the data do suggest that these microcharacters are potentially 
useful in Vanuatu as polyploid indicators. 

ANATOMY 
The species from Vanuatu show the basic vascular pattern described from other 
Tmesipteris species (Sykes 1908, Sahni 1925). Thus the solid core of tracheids in the 
rhizome becomes medullated and breaks up in the transition region to form a variable 
number of groups of tracheids arranged around a central pith. The representative 
sections illustrated in Fig. 3 show the typical arrangement of the stele at the top of the 
transition region. In all three species groups of tracheids can be seen surrounding a 
well defined pith. There are, however, differences in the nature of the pith cells and 
two basic types can be recognised. 



n= 104 


62.7 ± 4.2 


96.0 


n= 104 


57.5 ± 3.3 


92.5 


n= 104 


57.4 ± 3.8 


- 


n= 104 


62.0 ± 3.6 


93.2 


n = c.104 


58.6 ± 3.6 


92.3 


n = 208 


82.8 ± 4.1 


133.4 


n = c.208 


81.5 ± 4.3 


123.6 



TMESIPTERIS IN VANUATU 89 

+ *** ** z!m> * **. ** - v £ 



a 










* *£i?i* 



?? G fu R w p^r m . ane !l- acetoc f mine Preparations for meiosis. X 750. a) T. oblongifolia RSNH 
d™' %Stt£S2l£r m * "* bivalent* c) r van™* rsn'h 2354. ^Explana^ 



diagram showing 104 bivalents 



90 



FERN GAZETTE: VOLUME 13 PART 2 (1986) 





FIGURE 2. Permanent acetocarmine preparation of meiosis. X 750. a) T. oblanceolata RSNH 21 1 2. 
b) Explanatory diagram showing 208 bivalents. 



TMESIPTERIS IN VANUATU 



91 




^Mil'S^S^S'ST 8 ° f aeria ' Sh °° tS ° f Tmes 'P*e"s species from Vanuatu. X 1 00 a) T 
Sf»£pW^^ HSNH 21 1°2. l\ 



92 FERN GAZETTE: VOLUME 1 3 PART 2 (1 986) 



The first type is illustrated by T. oblongifolia (Fig. 3a) and T. vanuatensis (Fig. 3b). In 
these species the pith is small and made up of narrow thick walled, lignified cells 
which in transverse sections have the same appearance as fibres or sclerenchyma. 
The tracheids around the pith are by contrast thinner walled and the tracheid bundles 
can be located in the sections by their mesarch protoxylem. The second type of pith is 
characteristic of T oblanceolata (Fig. 3c,d). Here the groups of tracheids surround a 
relatively large pith made up of thin walled parenchymatous or slightly 
collenchymatous cells. Even in the distal parts of the leafy shoot, where the number of 
tracheid bundles and pith are much reduced, the pith cells are still essentially 
parenchymatous (Fig. 3d). Medullary xylem has been found associatedwiththistypeof 
pith in T. vieillardii (Sahni 1 925) from New Caledonia and T. oblanceolata (Braithwaite 
1973) from the Solomon Islands, but none has been found in the material from 
Vanuatu. On present evidence the pith type appears to be constant for each species. 

Variations in the pith cells in the stems of other Tmesipteris species have been 
reported by a number of previous investigators, notably Dangeard (1890-91), Sahni 
(1925) and Braithwaite (1973). In all cases two types of pith composed of either 
parenchymatous cells or sclerenchymatous cells have been recognised. The 
consistency of the observations so far suggests that this recently rather neglected 
anatomical character perhaps merits further investigation to establish whether it may 
be useful as a taxonomic indicator, either at the species level or for the grouping of 
species. 

ECOLOGY AND DISTRIBUTION 
During the 1971 Royal Society and Percy Sladen Expedition to the New Hebrides 
Tmesipteris was encountered only sporadically in montane forest at altitudes of 475- 
745m on Aneityum and Tanna in the south and 900- 1 650m on Espiritu Santo in the 
north. However, many of the other larger islands in the group attain altitudes in excess 
of 500m so that it can perhaps be anticipated that further collecting will extend its 
distribution within the islands. 

T. oblongifolia and T. vanuatensis were confined almost exclusively to the lower 
parts of the tree fern trunks belonging to the genus Cyathea. Both species were 
recorded from C. lunulata, but they were also collected from other Cyathea species so 
that they do not appear to be confined to any particular species. 

Unlike the other two species, T. oblanceolata was never found on tree ferns. It 
grew among mosses and filmy ferns on the trunks of angiospermous trees such as 
Metrosideros and Weinmannia (see Fig. 1b in Braithwaite 1975) or out of organic 
accumulations beneath epiphytic ferns such as Asplenium nidus. T. oblanceolata has 
also been recorded on decayed wood on the forest floor (Milne 272K). Non-tree-fern 
substrates have been reported for Tmesipteris species elsewhere; notably T. vieillardii 
of New Caledonia (Sahni 1925), T tannensis (Spreng.) Bernh. s. strict, in New Zealand 
(Chinnock 1976) and T. oblanceolata from the Solomon Islands (Braithwaite 1973). 

It is becoming increasingly clear that speciation within the genus has been 
accompanied by some ecological differentiation, although the extent to which the 
latter may be useful in delimiting species or determining species relationships is at 
present not clear. 



TMESIPTERIS IN VANUATU 93 



KEY TO THE SPECIES 
Aerial shoots 6- 16cm long, leaves ovate-oblong or narrowly oblong to elliptic. 

Leaves ovate-oblong with obtuse apices, 

l/b ratio <3, distichously arranged 

beyond the sporophylls. T. oblongifolia 

Leaves narrowly oblong to elliptic with 

acute apices, l/b ratio >3, spirally 

arranged beyond the sporophylls. T. vanuatensis 

Aerial shoots 15-38cm, leaves narrowly 

rectangular to narrowly obovate with 

truncate or rounded apices. T. oblanceolata 

Tmesipteris oblongifolia A. Braith. sp. nov. 

Planta epiphytica in truncis filicum arborum. Surculis aerius simplex, pendulus, (6-)8- 
14{-16)cm longus, per unum crescentem maturescens, folio magno terminati. Folia 
infra sporophylla spiraliter disposita, supra sporophylla disticha disposita, subcoriacea, 
(7-)9-13(-15)mm longa, (3-)3.5-4.5(-5)mm lata, ovato-oblonga, apicibus rotundatis 
obtusis mucronatis. Sporophylla spiraliter disposita, medianum caulis foliosi 
occupantes, longitudine folia aequantia. Synangium 2-4mm longum, sporangiis lobiis 
aequalibus. Sporae biiaterales, monoletae, concavo-convexae, (51-)57-62(-73)jjm 
longae, (18-)21-22(-25)um latae. Chromosomatum numerus gametophyticae 104. 

Holotype: Tanna, W ridge of Mt. Toukosmeru (1 9°33'S 1 69°21 'E), 500m, epiphyte on 
Cyathea trunk (same sp. as RSNH 2184), 28 Jul. 1971, A.F. Braithwaite RSNH 2204 
(K). 

Plant epiphytic on trunks of tree ferns. Aerial shoots simple, pendulous, (6-)8-14(- 
16)cm long, maturing in one growing season and terminating in large leaf-like 
appendage. Leaves spirally arranged below the sporophylls, distichously arranged in 
portion of leafy shoot distal to the sporophylis, 3-4 per cm stem, subcoriaceous, (7-)9- 
13(-15)mm long, (3-)3.5-4.5(-5)mm wide, ovate-oblong with rounded obtuse 
mucronate apex. Sporophylls spirally arranged, 5 per cm stem, occupying the middle of 
the leafy part of the shoot or throughout the upper two thirds, ± equal in length or 
slightly shorter than the leaves. Synangia 2-4mm long, 1-1. 5mm high, with lobes of 
sporangia approximately equal, ± globular. Spores bilateral, monolete, concavo- 
convex, (51-)57-62(-73)jjm long, (1 8-)21-22(-25)jjm broad. Chromosome number n = 
104. 

T. oblongifolia is closely allied to T. lanceolata from New Caledonia and New 
Zealand. The two species are similar in size and share a distichous arrangement of 
leaves in that portion of the leafy shoot distal to the sporophylls. The two species also 
possess the same pith type in the stem and chromosome number. Dangeard (1 890-91) 
describes and illustrates the pith cells of T. lanceolata as "fibres medullaires" and 
unpublished chromosome counts by the author show it to be a tetraploid. T. 
oblongifolia can however be distinguished by its thinner texture, ovate-oblong leaves 
with an obtuse apex and by the position of the sporophylls, which are found in the 
middle or upper two thirds of the leafy shoot and never only at the base or in the lower 
half as in T. lanceolata. 

Distribution: Philippines, Vanuatu and the Marquesas. 

Specimens examined: 

VANUATU. Aneityum, ridge leading to Woptiabo, c.5km ENE of Anelcauhat (20°13'S 

169°49'E), 487m, epiphytic on Cyathea lunulata in ridge side forest, 23 Jul. 1971, 

Braithwaite RSNH 21 46 (K). Tanna, W ridge of Mt. Toukosmeru (1 9°33'S 1 69°21 'E), 

644m, epiphytic on base of large Cyathea lunulata, 28 Jul. 1971, Braithwaite RSNH 

2211 (K). 

MARQUESAS. Feani, vieux sentrer 'Atuona a Hanamenu, haute valine cote 

Hanamenu, 850m, brousse fougeres arborescentes et Freycinetia, Epiphyte sur les 

bases de fougeres arborescentes, assay rare, 5 Mar. 1 975, Schafer & Oline 5272 (K); 



94 FERN GAZETTE: VOLUME 13 PART 2 (1986) 



Chemin d'Omoa a Hanavave, crete principale, mont Moratina (Mt. Boise de la carte), 
670m, brousse assay humide avecCyathea, Crossostylis, sur bases deCyathea, assay 
rare, 18 Sept. 1975, Schafer 5758 (K); Feani, montagnes entre la haute valine de 
Hanamenu et la crete de Temetiu, 900m, petit haut valine a foret tres humide: 
Crossostylis, Cyathea, Pandanaceae, Weinmannia, Myrsine, sur troncs de Cyathea. 
assay rare, 23 Oct. 1975, Schafer 5914 (K); Nukuhiva, Quayle 1305 (K). 
PHILIPPINES. Mindanao, Davao Dist: Mt. Apo 6000ft, epiphyte always on trunks of tree 
ferns, Oct. 1904, Copeland 1433 (K); Mt. Apo, May 1909, Elmer 10600 (BM, K); Mt. 
Apo, 9000ft, Feb. 1929, Hachisaka s.n. (BM): Mt. Apo, 1800m, 1932, Copeland 203 
(BM). 

Tmesipteris vanuatensis A. Braith. sp. nov. 

Planta epiphytica in truncis filicum arborum. Surculis aerius simplex, pendulus, (6-)8- 
1 5(- 1 6)cm longus, foliis et sporophyllis spiraliter dispositis et folio magno terminatis. 
Folia subcoriacea, 8-14mm longa, 2.5- 3. 5mm lata, anguste oblonga vel anguste 
elliptica, apicibus acutis et mucrone setaceo 0.5- 1.0mm longo. Sporophylla medianum 
caulis foliosi occupantes, longitudine folia subaequantia. Synangium parvum, 3- 
3.5mm longum, globosum, sporangiis lobis aequantibus. Sporae monoletae, concavo- 
convexae, (50-)59-62(-67)jjm longae, (18-)21(-25)um latae. Chromosomatum 
numerus gametophyticae 104. 

Holotype. Espiritu Santo, crest of NW ridge of Mt.Tabwemasana, c. 1 600m, epiphyte on 
Cyathea sp. in ridge top Metrosideros-Weinmannia forest, 2 Sept. 1971, A.F. 
Braithwaite RSNH 2382 (K). 

Plant epiphytic on the trunks of tree ferns. Aerial shoots simple, pendulous, (6-)8- 1 5(- 
16)cm long, maturing in one growing season, with leaves and sporophylls spirally 
arranged and terminating in a large leaf-like appendage. Leaves subcoriaceous, 8- 
14mm long, 2.5-3. 5mm, narrowly oblong or ovate-oblong to narrowly elliptical with 
acute apices and bristle-like mucro 0.5-1. Omm long. Sporophylls occupyingthe middle 
or throughout leafy part of shoot, equal to or slightly shorter than the leaves. Synangia 
3-3. 5mm long, globose, persistent, with two equal sporangial lobes. Spores monolete, 
concavo-convex, (50-)59-62(-67)pm long, (18-)21(-25)jjm broad. Chromosome 
number n = 104. 

T. vanuatensis can be distinguished from T. oblongifolia by its spirally arranged 
narrowly oblong to almost elliptical leaves with a larger length/breadth ratio and 
generally acute apices. The leaves are also more widely spaced and arise from the 
stem at a more acute angle giving the plant a generally more lax and slender 
appearance than T. oblongifolia (Fig. 4). 
Known only from Mt. Tabwemasana, Espiritu Santo. 

Other specimen examined: 

Espiritu Santo. Camp site no. 4, Nokovula Village, 23k SSWof Malau, Big Bay(1 5°20'S 
1 66°44'E), disturbed forest area below village, c. 900m, epiphyte on Cyathea lunu/ata, 
1 Sept. 1971, Braithwaite RSNH 2354 (K). 

Tmesipteris oblanceolata Copel., Philip. J. Sci. 60: 99(1 936); A. Braith., Brit. Fern Gaz. 
10: 296(1973). 

Type: Solomon Islands, Guadalcanal, Tutuve Mt, 1700m, Kajewski 2632 (A). 

Plants epiphytic on angiospermous tree trunks or growing on moss covered decaying 
wood on forest floor. Aerial shoots simple, pendulous or occasionally sub-erect, 
(15-)20-30(-38)cm long, maturing in a single growing season and terminated by a 
small leaf-like appendage; transition region (4-)5-10(-1 1)cm long; leaves and 
sporophylls spirally arranged and often tending to decrease in size towards the apex. 
Leaves (10-)1 1-14(-16)mm long, 2. 5-3. 5mm broad, coriaceous to subcoriaceous, 
(4-)5(-6) per cm stem, narrowly oblong or rectangular but narrowing towards the base 
to narrowly obovate, ± falcate, apex truncate and sometimes bilobed to rounded, 
mucronate; mucro stiff, 1-2mm long. Sporophylls developed in mid region or 
throughout upper two thirds of leafy shoot, equal in length to leaves; synangia 3-4mm 
long, 1.5-2mm high, persistent, bilocular, sporangial lobes ± equal. Spores monolete, 
concavo-convex, (73-)81-83(-94)^im long, (25-)31(-36)um broad. Chromosome 
number n = 208. 



TMESIPTERIS IN VANUATU 



95 



\ \ 



■ MM 



I 




• 



V 



\ 



a 



FIGURE 4. a) T. oblongifolia RSNH 2204. b) T. vanuatensis RSNH 2382. Specimens preserved in 
alcohol. X 1/2. 



T. oblanceolata in Vanuatu is rather variable with respect to length of the aerial 
shoot and in size and shape of the leaves. The plants from Espiritu Santo with aerial 
shoots up to 26cm long bearing short narrowly obovate leaves with rounded apices, 
are virtually indistinguishable from T. oblanceolata Copel. from the Solomon Islands, 
except that medullary xylem found in specimens from the Solomon Islands 
(Braithwaite 1973) was not found in the single specimen available for sectioning from 
Espiritu Santo. The pith is otherwise of a similar parenchymatous type. The more 
extensive collections from Aneityum have aerial shoots up to 38cm long, some 
possessing narrow slightly obovate leaves with rounded apices, while the majority 
have longer, narrowly rectangular leaves with truncate apices. Similar variation is 
evident in material attributed here to T. oblanceolata from New Caledonia and Samoa. 

The plants from Aneityum, with leaves with markedly truncate sometimes almost 
bilobed apices, are very similar to the Australian species T. truncata (R.Br.) Desv. They 
also share the same chromosome number. Barber (1957) reported a chromosome 
number of n = 201-21 1 for T. truncata from several different localities in New South 
Wales and n = 208 has been found here in material from Aneityum. There are however 
some ecological and anatomical differences. The plants from Aneityum are either 
epiphytic on angiospermous trees or terrestrial on decayed wood while T. truncata 
from Australia is generally, though not exclusively, epiphytic on tree ferns. There are 
two specimens from Australia at Kew collected from non-tree-fern substrates; one 
with typical truncate leaves labelled "Macquarie Harbour, humid rocky banks B 
mountains in shaded woods, 1825, Cunningham 92"; and the other with more 
oblanceolate leaves labelled "Head of Clyde River, 25 miles SSWof Howra, 2000ft, on 
mossy ledge of sandstone cliff, 2 May 1937, Rodway 2369". The anatomical difference 
concerns the pith type in the stem. Dangeard (1 890-91) reported a sclerenchymatous 



96 FERN GAZETTE: VOLUME 1 3 PART 2 (1 986) 



pith type in T. truncata from Australia which differs from the parenchymatous pith 
found here in the material from Vanuatu. However Dangeard (in contrast to the often 
very detailed drawings from the stems of the other species he examined) shows only a 
very small outline tissue sketch for the stem of T. truncata (Plate XIV, Fig. 1 1) and it 
would be desirable to have the detail confirmed for material both from tree-fern trunks 
and from non-tree-fern substrates. It is possible that variation in the pith type may be 
correlated with differences in ecology. 

T. oblanceolata and T. truncata are clearly very closely related and they probably 
represent forms of the same species. However on present evidence it is difficult to 
assess the extent or taxonomic significance of their ecological and anatomical 
differences. Future studies may well confirm their conspecificity but it is considered 
preferable for the time being to segregate the Pacific material under T. oblanceolata. 

Distribution: New Caledonia, Vanuatu, Solomon Islands, Fiji, Samoa. 

Specimens examined: 

VANUATU. Aneityum: ridge crest N of Woptiabo, S end of Nithuon Nelvau (20°13'S 

169°49'E), 640m, ridge top forest with Metrosideros, epiphytic on large leaning tree on 

ridge top growing underneath a large plant of Asplenium nidus, 23 Jul. 1971, 

Braithwaite RSNH 2152(K); c. 5km NE by N of Anelcauhaut, on crest of ridge running S 

from Inrero (20°11'S 1 69°47'E), 745m, epiphytic on trunk of Metrosideros, 

occasionally on forest floor, 21 Jul. 1971, Braithwaite RSNH 2112(K); high grounds, 

decayed trees, Nov. 1 853, Milne 272 (K); 1 854, Seeman s.n. (BM); crSte S de I'lnrero, 

alt. 750m, epiphyte au bas de troncs, fronde portee ± horizontalement, 23 Jul. 1971, 

Raynal RSNH 1 61 47 (K). Espiritu Santo: northern ridge of Mt. Tabwemasana (1 5°22'S 

166°45'E), 1650m, low forest on ridge crest, Weinmannia dominant, 4 Sept. 1971 

Raynal RSNH 16386 (P). 

SOLOMON ISLANDS. Guadalcanal, Mt. Popomanaseu, halfway between upper camp 

and Vunuvelakama, c. 5000ft, growing erect in moss around base of trees in ridge top 

moss forest, 3 Nov. 1965, Braithwaite RSS 4782 (K). 

NEW CALEDONIA. Mt. Koghi, kauri ft. 1 000ft, on prostrate decaying trunk covered with 

liverworts, 13 Jun. 1914, Compton 764 (BM). 

LORD HOWE ISLAND. Epiphyte on the top of Mt. Gower in mist forest, 2600ft, Aug. 

1965, Game 65/1 /SN (K). 

SAMOA. May 1876, Whitmee s.n. (K). 

ACKNOWLEDGEMENTS 
The author expresses his gratitude tothe Royal Society of London for the opportunity to 
participate in the Royal Society and Percy Sladen Expedition to the New Hebrides 1971 
and gratefully acknowledges the photographic assistance of Mr B.V. Case during 
preparation of the illustrations. 

REFERENCES 
BARBER, H.N. 1957. Polyploidy in the Psilotales. Proc. Lin. Soc. N.S.W. 82: 201-208. 
BERNHARDI, J.J. 1801. Tentamen alterum filices in genera redigendi. Schrad. J. Bot. 1800(2): 

131, t.2, f.5. 
BRAITHWAITE, A.F. 1973. Tmesipteris in the Solomon Islands. Brit. Fern Gaz. 10: 293-303. 
BRAITHWAITE, A.F. 1 975. The phytogeographical relationships and origin of the New Hebrides 

fern flora. Phil. Trans. R. Soc. Lond. B. 272: 293-313. 
CHINNOCK, R.J. 1976. The identification, typification and origin of Tmesipteris tannensis 

(Psilotaceae). Taxon. 25: 115-121. 
DANGEARD, P. A. 1890-91. Memoire sur la morphologie & I'anatomie des Tmesipteris. Le 

Botaniste, Series II: 163-222. 
LOVIS, J.D. 1977. Evolutionary patterns and processes in ferns. Adv. Bot. Res. 4: 229-419. 
MANTON, I. 1 950. Problems of cytology and evolution in the Pteridophyta. Cambridge University 

Press. 
SAHNI, B. 1925. On Tmesipteris vieillardii Dangeard, an erect terrestrial species from New 

Caledonia. Phil. Trans Roy. Soc. B. 213: 143-170. 
SPRENGEL, K. 1800. Bemerkungen uber einige kryptogamische Pflanzen. Schrad. J. Bot. 

1799(2): 267. 
SYKES, M.G. 1908. The anatomy and morphology of Tmesipteris. Ann. Bot. 22: 63-89. 



FERN GAZ. 13(2) 1986 97 



EFFECTS OF SALINITY ON GAMETOPHYTE GROWTH OF 

ACROSTICHUM AUREUM AND A. DANAEIFOLIUM 



ROBERT M. LLOYD & DONALD P. BUCKLEY 
Department of Botany, Ohio University, Athens, Ohio 45701, U.S.A. 

ABSTRACT 

Spore germination and gametophyte growth under salinity regimes varying from 
O.Oto 3.0% NaCI was studied to determine the stress tolerance of the gametophyte 
generation. Responses of both New World species of Acrostichum are similar to 
those of other mangrove species which have been studied. A. aureum has slightly 
greater tolerance to increased salinity than A danaeifolium. Growth responses of 
A. aureum suggest it can be classified as a true halophyte, whereas those of A 
danaeifolium suggest it is a semi-halophyte. The response of the gametophyte 
generation of these species to salinity parallels the natural habitats of the 
sporophyte generation. 



INTRODUCTION 
Halophytes are notably rare in pteridophytes. The most well known example is the 
mangrove fern genus Acrostichum. There are three species circumscribed in this 
genus, each with different apparent tolerance to salinity. The observed habitat 
variation of the species forms a continuum from fresh water to inundation bytides and 
sea water. The most widely distributed species is A. aureum L., circumtropical in 
distribution and frequently forming large colonies in mangrove swamps, salt- and 
brackish-marshes, and low hammocks near sea water (Holttum 1955; Small 1938; 
Walsh 1 974). The remaining two species are much more restricted in distribution. In 
tropical Asia and Australia, A. speciosum Willd. occurs in mangroves (Walsh 1974) 
frequently inundated by tides and has a greater tolerance for sea water than A. aureum 
(Holttum 1 955). Acrostichum danaeifolium Langsd. & Fisch. is distributed in the New 
World tropics and subtropics in fresh or brackish water swamps, lakes, and ditches and 
along canal margins (Adams and Tomlinson 1 979). This species is frequently found 
inland from coastal regions, sometimes associated with pines and palms or other 
glycophytes. 

One oftheprimaryfactorsdeterminingthe growth anddistribution of plants in salt 
marsh habitats is the level of soil salinity (Jefferies et al. 1 979). Other factors include 
both intraspecific and interspecific competition, especially when species may be only 
facultative halophytes. Barbour (1 970) has questioned whether all halophytes are just 
facultative halophytes and has suggested that the ability to reproduce under 
"halophytic" conditions should bethe ultimatecriterion of salttolerance. In halophytic 
pteridophytes, for sexual reproduction to be successful the gametophytic and 
sporophytic generation must succeed. Therefore, both generations should exhibit 
parallel tolerances to stress under soil salinity, assuming that the soil salinity 
conditions of the gametophytic and sporophytic habitats parallel one another. 
However, Ungar (1978) has reportedthat surface soils may havesalinitiesfromtwoto 
1 00 times that of subsoils. Thus, the gametophytic stage may be critical if a species is 
to successfully inhabit a saline environment. 

In the New World, collections have been made of both Acrostichum species. 
Gametophytes originating from spores of these plants as well as from the fresh water 
aquatic, Ceratopteris thalictroides (L.) Tod., have been grown under a variety of salinity 
conditions to test the hypothesis that tolerance to salinity by the gametophytic 
generation will parallel the habitat conditions in which the sporophytes occur. 



98 



FERN GAZETTE: VOLUME 13 PART 2 (1986) 



MATERIALS AND METHODS 
Spores utilized in this study were collected from the following locations:/!, aureum: 
culture 190, Jamaica, Westmoreland Parish, 0.25 mile east of Negril on road to 
Savana la Mar. Plants occur densely in a large population in a lowland coastal swamp 
which is periodically inundated by tides; culture 1 93, Panama (Canal Zone), very large 
population of over 3000 individuals in mangrove swamp about 0.2 mile from road to 
Colon on road to Coco Solo; culture 150, Florida, Dade Co., 30.5 miles southwest of 
entrance of Everglades National Park, at road to Westlake, growing in Rhizophora- 
swamp; A. danaeifolium: culture 204, Florida, open marsh with about 40 individuals, 
Collier Co., 0.7 mile south of state route 92 on U.S. Highway 41, about 3.5 miles inland 
from coast; Ceratopteris thalictroides: culture 174, Guyana, two miles east of 
Georgetown on main coastal public road, in wet marsh with Nymphaea next to 
gasoline station; culture Hawaii, taro patches, Hawaiian Islands, inundated weekly 
with fresh water. Adscript numbers and letters designate spore progenies from 
different individual sporophytes collected at each location. 

Spores were sown and gametophytes grown on inorganic nutrient medium 
solidified with 1% agar (see Klekowski 1969, for composition) in 100x 15mm petri 
dishes under continuous illumination from fluorescent and incandescent lamps at 
about 23°C. Nutrient mediaweresupplementedwith NaClpriortothe addition of agar, 
yielding concentrations of 0.5 to 3.0%. NaCI concentration was ascertained by 
conductivity measurements (from 0.8mmhos/cm at 25°C in control to 44mmhos/cm 
in 3.0% NaCI.) Conductivity was measured with a Radiometer CDM2 conductivity 
meter. Data for 190-D/193-B (Table 4) represent gametophytes of both progenies 
transferred on to single petri dishes. 



TABLE 1 
Percent spore germination \n Acrostichum (1 5 days following sowing) and Ceratopteris (12 days 
following sowing) at salinity regimes varying from 0.0% to 3.0% NaCI. (Sample size = 100). 

Species and Culture Number 





A. aureum 






A. dan a 


eifolium 


C. thalictroides 




NaCI 


190-D 


193-B 


193-K 


204-2 


204-8 


174- A 


Hawaii- 


Hawaii- 


(%) 














A 


B 


0.0 


69(100*) 


66(96) 


76(100) 


99 


89(100) 


86(96) 


95(100) 


90(100) 


0.5 


55(80) 


69(100) 


61(80) 


99 


67(75) 


90(100) 


50(52) 


20(22) 


1.0 


51(74) 


60(87) 


47(62) 


92 


72(81) 


85(94) 


30(32) 





1.25 


1 9(28) 


44(64) 


54(71) 


72 


54(61) 


84(93) 








1.5 


11(16) 


43(62) 


56(74) 


24 


21(24) 


19(21) 








1.75 


3(4) 


19(28) 


17(22) 


28 


8(9) 


11(12) 








2.0 


1(14) 


8(12) 


17(22) 


18 


KD 


2(2.2) 








2.25 


2(3) 


2(3) 


5(7) 


3 














2.5 





1(14) 


4(5) 

















2.75 








1(13) 

















3.0 





1(1.4) 


1(1.3) 


















'Values given in () are adjusted to 100% to adjust for variation in intersporophytic spore viability. 



SALINITY EFFECTS ON ACROSTICHUM GAMETOPHYTES 99 



RESULTS 
Spore germination in all three species initially occurred five to six days following 
sowing and was highest in the control andthe lowest NaCI concentration. There was a 
differential response of spore progenies from individual sporophytes of each species to 
each of the culture regimes. In Acrostichum, germination rates differ slightly at the 
higher NaCI concentrations (Table 1). Spore progenies of both species exhibited 
consistent but decreasing levels of germination upto 1.25% NaCI. At salinities above 
1.25 and 1.5%, respectively, in A danaeifolium and/4, aureum, there are significant 
decreases in germination. At NaCI concentrations between 2.25 and 3.0%, spore 
progenies of A. aureum exhibited low levels of germination, whereas in the other 
species germination did not occur. There are indications of bimodality (stress pulses) in 
three of the five dose response curves (193-K, 204-2, 204-8 at salinities of 1.25-1.5%, 
1.75%, and 1.0% respectively). If thispattern isreal itcouldresultfromtheoperationof 
two physiological processes whose reaction optima occur at different levels of salinity. 

In Ceratopteris, germination was greatly inhibited in Hawaiian spore progenies at 
all NaCI concentrations and did not occur above 1.0%. The Guyana sample is of 
potential interest, however, as germination responses were similar to those of A. 
danaeifolium. 

The fraction of gametophytes to attain two-dimensional growth within 1 5 days of 
sowing was determined in order to evaluate the effect of salinity on developmental 
rates (Table 2). There is no appreciable difference in the two species of Acrostichum at 
the lower salinities with almost all gametophytes reaching thetwo-dimensional stage. 
The reaction curves for developmental rate versus salinity are markedly bimodal for 
the spore progenies of A. aureum tested. Developmental rates decreased 
synchronously for all four samples from both species between 1.0 and 1.25% NaCI. 
The second range of salinities (1.25 to 1.75%) which resulted in rapid gametophyte 
development for A. aureum produced reduced development in culture 204-2 and 
severely inhibited development at 1.75% NaCI in culture 204-8of A danaeifolium. The 
phenomenon responsible for the higher developmental rate in higher salinities for A 
aureum appears to extend this species' maximum salinity tolerance beyond that of A 
danaeifolium. However, it was not determined what proportion of 15 day old one- 
dimensional gametophytes later attained the two-dimensional stage under 
hypersalinity conditions. 

TABLE 2 
Mean percent attainment of two-dimensional morphology in gametophytes of Acrostichum 
grown under varying salinity regimes 15 days following sowing. 

Species and Culture Number 





A. aureum 




A. danaeifolium 


NaCI (%) 


193-B 


193-K 


204-2 


204 


0.0 


100 


97 


100 


100 


0.5 


100 


96 


100 


100 


1.0 


100 


97 


97 


98 


1.25 


90 


84 


85 


93 


1.5 


97 


90 


80 


87 


1.75 


97 


86 


83 


14 


2.0 


75 


54 








2.25 















Sample size variable and dependent upon number of available gametophytes: 30-45 in 0.0% to 
1.75%; (1)10-29 in 2.00%; 11-14 in 2.25%; less than 5 in remaining regimes. 



100 



FERN GAZETTE: VOLUME 13 PART 2 (1986) 



TABLE 3 

Maximum and (mean) gametophyte size (in sq. mm) attained in cultures oiAcrostichum at salinity 

regimes varying from 0.0% to 2.25%. Sample size = 10. 

Species and Culture Number 

A. danaeifolium 



0.0 

0.5 

1.0 

1.25 

1,5 

1.75 

2.0 

2.25 



A. aureum 



NaCI (%) 190-D 



2.8(1.4) 
1.9(1.0) 
2.0(0.9) 
0.4(0.3) 
1.1(0.6) 



193-B* 

10.9(3.8) 
5.9(2.8) 
4.0(1.8) 
2.5(1.2) 
1.2(0.7) 
0.6(0.4) 
0.1(0.1) 
0.2(0.1) 



193-K** 

4.1(2.4) 
13.9(2.0) 
4.5(2.0) 
1.8(0.8) 
0.4(0.2) 
0.4(0.4) 
0.2(0.1) 
0.09(0.07) 



204-2*** 

11.8(5.1) 
6.0(3.1) 
4.0(2.1) 
2.4(1.2) 
0.7(0.4) 



204-8*** 

6.3(4.3) 
8.5(3.2) 
2.8(1.2) 
1.9(0.9) 
0.8(0.4) 
0.09(0.08) 



sampled 19 days following sowing, 
sampled 18 days following sowing, 
sampled 21 days following sowing. 



TABLE 4 

Mean percent non-chlorotic tissue in gametophytes of A. aureum grown 70 days in nutrient 

control medium and then transferred to variable salinity regimes. 



NaCI (%) 

0.0 

1.0 

1.25 

1.75 

2.0 

2.25 

2.5 

2.75 

3.0 



150-B: Days from transfer 



190-D /193-B: Days from transfer 



14 



21 



4 



15 



99 


91 


91 


100 


100 


100 


00 


99 


99 


100 


100 


100 


94 


86 


94 


100 


99 


92 


91 


80 


83 


97 


91 


69 


71 


74 


75 


97 


90 


72 


73 


62 


58 


87 


74 


69 


40 


27 





76 


81 


39 


45 


31 





38 


25 


13 


40 


30 





32 


23 






Maximum and mean gametophyte area was measured 18 to 21 days following 
sowing to evaluate the effect of salinity on growth rates (Table 3). The growth rate of/4. 
danaeifolium appears to be greatest in the 0% NaCI controls and then decreases 
linearly with increasing levels of salinity. Low NaCI concentrations (0.5 and 1.0%) 
reduced growth rates of A. danaeifolium to a greater extent than those of A aureum. 
For example, reduction of mean gametophyte size (compared to 0.0% NaCI) of A. 
aureum progenies varies from 1 6. 7 to 52. 7%, whereas reduction in size of A. danaei- 
folium progenies is 58.9 to 72.1%. 

Salt stress and development of chlorotic tissue was measured in gametophytes of 
A. aureum (Table 4). These gametophytes were grown to maturity for 70 days on 
control media and then transferred to various salinity regimes. In culture 1 50B, below 
2.0% NaCI there is no apparent difference over time but only with salinity 
concentration. The effect appears to be initial and then persistent. There is some 
indication of a stress pulse at 1.25 to 2.0% NaCI with gametophytic tissue recovery 
after 21 days. Above 2.25%, the salinity effects are progressive over time. In culture 
1 90/ 1 93, effects appear to be absent below 1 . 25% NaCI. At levels of 1 . 25% and above, 
there is a progressive increase in chlorotic gametophytic tissue with both time and 
NaCI concentration. 



SALINITY EFFECTS ON ACROSTICHUM GAMETOPHYTES 101 



DISCUSSION 
The tolerance to salinity of gametophytes of both species of Acrostichum appears to be 
significantly greater than two of the three progenies of the glycophyte Ceratopteris 
thalictroides. Warne and Hickok (pers. comm.) have also studied NaCI effects on spore 
progenies of C. thalictroides. They analyzed spore germination and gametophyte 
survival of progenies from ten sporophytes from various parts of the World. These 
progenies exhibit a wide range of tolerance to NaCI. At 0.8% NaCI, spore germination 
in six of the ten progenies was reduced by 4.0 to 15.9 (mean = 7. 53%) compared to the 
controls (0.0% NaCI). In the remaining four progenies, germination was reduced 48.8 
to 97.3 (mean = 77.7)%. Twenty-one days following sowing, gametophytes of the 
former group were 2.5 to 10% the size of control gametophytes, whereas 
gametophytes of the latter group failed to survive. These results indicate that some 
populations of Ceratopteris may have salinity tolerance approaching that exhibited by 
A danaeifolium. Recently, Petersen (1985) reported that spores of A. danaeifolium 
were capable of germinating in up to 2.5% NaCI whereas spores of the glycophytic 
species Osmunda spp. and Onoclea sensibilis ceased germination at 0.6% NaCI. 

In Acrostichum, there is a wide range of tolerance to salinity, although A aureum 
consistently shows a slightly greater tolerance at slightly higher NaCI levels. In spore 
germination, two of the three progeny samples of A. aureum show 50% germination at 
NaCI concentrations of 1.5 and 1.75%. In A. danaeifolium, 50% germination occurs 
between 1.25 and 1.5%. Similarly, the critical salinity level for attainment of two- 
dimensional morphology is 1.75 to 2.0% NaCI in A. aureum and 1.5 to 1.75% in A. 
danaeifolium. In addition, some progenies of mature gametophytes of A aureum can 
tolerate prolonged exposure to salinities of 2.0 to 2.25%. These results suggest that 
the critical soil salinity which will limit gametophyte survival of A danaeifolium will be 
about 1.5% and of some plants of/4, aureum, about 1.75 to 2.0%. However, due to the 
extremely large number of spores produced by individual sporophytes of these species, 
even very low percentages of survival at higher salinities could result in millions of 
successful gametophytes. 

Germination and growth in Acrostichum parallels that of other mangrove species 
which have been studied and have optimal growth between 0.6 and 1.5% NaCI 
(Connor 1969; Pannier 1959; Patil 1964). 

The spore germination and gametophyte growth patterns of A. danaeifolium are 
similar to those described for semi-halophytes by Waisel (1972). Semi-halophytes 
show a slow decrease in growth at initial stages of salinity increase, followed by a 
steady decrease with increasing salinity. On the other hand, in A aureum, the stress- 
pulse growth phases followed by a steady decrease in growth with increasing 
salinities is similar to the pattern of true halophytes. These studies indicate, therefore, 
that species of Acrostichum can be considered to be semi-halophytes or halophytes 
and that the salinity tolerance by the gametophyte generation is an integral part of 
their life-history. 

ACKNOWLEDGEMENTS 
The technical help of I. A. Ungar and T.R. Warne is gratefully acknowledged. This 
research was supported by grant number BMS- 75071 91 from the National Science 
Foundation. 

REFERENCES 
ADAMS, DC. & TOMLINSON, P.B. 1979. Acrostichum in Florida. Amer. Fern J. 69: 42-46. 
BARBOUR, M.G. 1970. Is any angiosperm an obligate halophyte?4A77e/-. Midi. Nat. 84: 105-120. 
CONNOR, D.J. 1969. Growth of grey mangrove (Avicennia marina) in nutrient culture. Biotropica 
1: 36-40. 



102 FERN GAZETTE: VOLUME 13 PART 2 (1986) 



HOLTTUM, R.E. 1 955. Flora of Malaya, Vol. II, Ferns of Malaya. Gov't. Printing Office, Singapore. 
JEFFERIES, R.L., DAVY, A.J. & RUDMIK, T. 1 979. The growth strategies of coastal halophytes. In 

R.L. Jefferies & A.J. Davy (eds.) Ecological Processes in Coastal Environments. Blackwell 

Scientific Publ., Oxford, 243-268. 
KLEKOWSKI, E.J. JR. 1969. Reproductive biology of the Pteridophyta. III. A study of the 

Blechnaceae. Bot. J. Linn. Soc. 62: 361-377. 
PANNIER, P. 1959. El efecto de distintas concentrationes sotenas sobre el desarrolo de 

Rhizophora mangle L. Acta cient. Venez. 10: 68-78. 
PATIL, R.P. 1964. Cultivation of mangrove seedlings in pots at Allahabad. U. P. Sci. Cult. 30: 43- 

44. 
PETERSEN, R.L. 1 985. Use of fern spores and gametophytes in toxicity assessments. Proc. Roy. 

Acad. Edin. 86B: 453. 
SMALL, J.K. 1 938. Ferns of the Southeastern States. Facsimile Ed. (1 964), Hafner Publ. Co., New 

York. 
UNGAR, I. A. 1978. Halophyte seed germination. Bot. Rev. 44: 233-264. 
WAISEL, Y. 1972. The Biology of Halophytes. Academic Press, New York and London. 
WALSH, G.E. 1974. Mangroves: A review. In R.J. Reimold & W.H. Queen (eds.), Ecology of 

Halophytes, Academic Press, New York and London. 



FERN GAZ. 13(2) 1986 103 



THE ECOLOGY OF PTERIDOPHYTES IN THE MWANIHANA 
FOREST RESERVE, TANZANIA 



J.C. LOVETT and D.W. THOMAS 
Missouri Botanical Garden, P.O. Box 299, St Louis, Missouri, 63166, U.S.A. 

ABSTRACT 

A collection of pteridophytes made in the Mwanihana Forest Reserve, Tanzania is 
described and related to the ecology of the forest. In all 78 species were collected, 
demonstrating the richness of the area in comparison with the total number of 500 
species estimated for the whole of tropical Africa. 

INTRODUCTION 
The Mwanihana Forest Reserve (7°50'S 36°55'E) is located on the steep east facing 
escarpment of the Uzungwa Mts, overlooking the Kilombero Valley and Selous Game 
Reserve. These mountains are composed of Pre-Cambrian crystalline gneiss, and have 
probably been in existence since the Cretaceous. The rainfall is relatively high for 
Tanzania, being estimated to be 2, 000- 2, 500mm a year with one wet season receiving 
greater than 1 00mm of rain a month between November and May on average, which 
has a peak in March and April (DHV, 1982). The dry season, between July and October, 
receives less than 50mm of monthly rain on average, with months of no rain common. 

Despite the marked dry season the escarpment is covered by continuous closed 
moist forest from an altitude of 450m to 1,800m, and in the past extended into the 
Kilombero flood plain where it has now been replaced by sugar plantation. These 
forests are part of the Eastern Arc group (Lovett, in press) which are notable for the 
high degree of endemism they contain, and it is now hopedthatthe Mwanihana Forest 
Reserve will become a National Park in order to protect many rare plants and animals. 

During the course of surveying the Reserve as part of the 1984 Uzungwa 
expedition supported by the National Geographic Society and World Wildlife Fund, a 
substantial collection of plants was made. This collection included some 78species of 
pteridophytes from the Mwanihana and nearby forest reserves which are presented 
here divided into various ecological categories, and as a check-list. The collections 
made were far from exhaustive, and there are probably something in the region of 100 
species of pteridophyte in the area. This is high for Tropical Africa, which is estimated 
to contain only 500 species altogether (Parris, 1985). Thus the small area of the 
Mwanihana Forest Reserve may contain as much as 20% of the total Tropical African 
pteriodophyte flora. 

Despite the incompleteness of the collections and the brief nature of the 
ecological observations it was thought necessary to publish them in order to draw 
attention to the high species richness of the Mwanihana Forest Reserve in the hope 
that further work may be stimulated. 

ECOLOGY 
The vegetation of Africa has been divided into a number of phytochoria based on plant 
species distribution and physiognomic types which are described by White (1983). 
Although the broad scope of this work renders it inaccurate at the small scale, it is 
useful as a general pattern within which to designate arbitrary ecological boundaries. 
Five types of forest can be recognised in the Mwanihana Forest Reserve according to 
this system, and they are briefly described below for the forest reserve using White's 
terminology. Ten examples of tree species which occur in each forest type in the 
reserve are also given. 



104 F ERN ! GAZETTE: VOLUME 13 PART 2 (1986) 



Within each forest type the pteridophyte habitat is divided into terrestrial, 
lithophytic, and epiphytic. Additional notes on the habitat follow the species name in 
parenthesis when necessary. All pteridophytes were growing in forest shade unless 
noted. These notes are self explanatory except for the following: an exposed habitat is 
one in partial shade; a stream habitat is on rocks in streams and river beds flowing 
through the forest. A "C" in parenthesis indicates that the species was collected at 
Chita (8° 30' S 35° 55' E), a locality to the south of the reserve, but also in the Uzungwa 
mountains. All other species were collected in the reserve. 

Zanzibar-lnhambane Lowland Forest 

Altitudinal range 450-750m, canopy height 25-30m with emergents to 40m. Large 

trees include: Afrosersalisia cerasifera; Albizia adianthifolia; Aningeria pseudo- 

racemosa; Chlorophora excelsa; Dialium holtzii; Erythrophleum suaveo/ens; Funtumia 

africana; Lettowianthus stellatus; Newton/a paucijuga; Terminalia sambesica. 

Terrestrial: Bolbitis acrostichoides, (C)*; Christella hispidula, (C); Microlepia 

speluncae; Pellaea doniana, (exposed). 

Lithophytic: Adiantum capi/lus-veneris, (exposed); Bolbitis sp. aff. acrostichoides, (C); 

Nephrolepis bis errata, (C). 

Zanzibar-lnhambane Intermediate Forest 

Altitudinal range 750- 1,200m, canopy height 25-30m with emergents to 40m. Large 
trees include; Anisophylla obtusifolia; Cassia angolensis; Cephalosphaera 
usambarensis; Ochna holstii; Octoknema orientalis; Sibangea pleioneura; Syzygium 
guineense; Tabernaemontana holstii; Trichilia dregeana; Uapaca pallidosa. 
Terrestrial: Amphineuron opulentum; Asplenium blastophorum; A. obscurum, (C); 
Bolbitis auriculata; B. gemmifera; Christella gueintziana, (C) (swamp); Lonchitis 
occidentalis; Pteris sp. aff. mildbraedii; P. sp. aff. prolifera; P. quadriaurita subsp. 
friesii; Tectaria gemmifera. 

Lithophytic: Antrophyum mannianum, (C) (stream); Asplenium formosum, (stream);A 
inaequilaterale; A. unilaterale; Bolbitis sp. aff. acrostichoides; Christella sp. aff. 
gueintziana, (stream); Elaphoglossum spathulatum, (stream); Menisorus paucif/orus, 
(stream); Sphaerostephanos arbuscula subsp. africanus, (stream). 
Epiphytic: Lycopodium phlegmaria, (canopy). 

Afromontane Rain-Forest 

Altitudinal range 1,200- 1,700m, canopy height 25-30m, with emergents to 35m. 
Large trees include: Allanblackia stuh/mannii; Bei/schmiedia kweo; Cassipourea 
gummif/ua; Cleistanthus polystachyus; Chrysophyllum gorungosanum; Myrianthus 
holstii; Newtonia buchananii; Ocotea kenyensis; Parinari excelsa; Strombosia 
scheff/eri. 

Terrestrial: Asplenium christii; A. sp. aff. gi/pinae; A. hypome/as; A. monanthes; A. 
Pvo/kensii; Athyrium scandicinum; Blechnum attenuatum; B. ivohibense; Blot iella 
natalensis; Ctenitis languinosa; Cyathea humilis, (ravine); Cyathea mossambicensis; 
Didymochlaena trunculata; Diplazium nemorale; D. pseudoporrectum; Dryopteris 
inaequalis; D. kifemens/s; Histiopteris incisa; Microlepia fadenii; Po/ystichum 
zambesiacum; Pneumatopteris usambarensis; Pteris buchananii; P. preussii; P. 
pteridioides; Tectaria gemmifera; Trichomanes cupressiodes; T. giganteum. 
Lithophytic: Asplenium boltonii, (damp, exposed); Elaphoglossum phanerophlebium; 
Hymenophyllum splendidum; Pellaea angulosa; Trichomanes borbonicum. 
Epiphytic: Asplenium aethiopicum; A. dregeanum; A. rutifolium; Be/visia spicata; 
Elaphoglossum acrostichoides; Hymenophyllum po/yanthos var. kuhnii; H. sib- 
thorpioides; H. splendidum; Lomariopsis warneckei; Lycopodium vertici/latum; 
Trichomanes giganteum. 

* = collected at Chita (see text above). 



ECOLOGY OF PTERIDOPHYTES IN MWANIHANA, TANZANIA 105 



Afromontane Undifferentiated Forest 

Altitudinal range 1,200- 1,800m, canopy height 15-25m. This forest type represents a 
drier or more exposed type of Afromonate Rain-Forest. Large trees include: Aphloia 
theriformis; Bequaetiodendron magalismontanum; Cryptocaria liebertiana; Ficalhoa 
laurifolia; Hirtella megacarpa; /sober/in/a scheffleri; Psydrax vulgar e subsp. 
rubrocristatum; Rapanea melanophloeos; Xylopia aethiopica; Xymalos monospora. 
Terrestrial: Schizaea dichotoma; Dicranopteris linearis, (exposed). 

Afromontane evergreen bushland and thicket 

Altitudinal range 1,700- 1,800m, canopy height 5- 10m. In the reserve this forest type 
is better referred to as Elfin Forest, and has been described for the nearby Uluguru 
mountains by Pocs (1976). It is a variant of Afromontane undifferentiated forest but is 
distinguished by the low canopy and rich cover of epiphytic bryophytes, indicating that 
a great deal of moisture must come from mist. Large trees include: Allanblackia 
ulugurensis; Apodytes dimidiata; Faurea saligna; Maytenus acuminata; Ocotea 
usambarensis; Olinia rochetiana; Podocarpus latifolius; Syzygium cordatum; 
Ternstroemia polypetala; Trichocladus goetzei. 

Terrestrial: Blechnum punctulatum. 

Lithophytic: Elaphoglossum macropodium. 

Epiphytic: Ctenopteris sp. aff. villosissima; Elaphoglossum macropodium; Grammitis 

kyimbilensis; G. nanodes; Lycopodium ophioglossoides; L dacrydioides; Pleopeltis 

excavata; Xiphopteris strange an a. 

Collection List 

The following is a list of the pteridophyte collections made in the Mwanihana Forest 
Reserve and Chita forests of the Uzungwa scarp. The numbers are those of D.W. 
Thomas' collections. The collection is at MO, with duplicates at DSM and K. All the 
species mentioned in this list are also cited in the ecology section of this paper. 

Pteridophyta 

Filicopsida 

Adiantaceae 

Adiantum capillus-veneris L., 3933 
Pellaea angulosa (Bory ex Willd.) Bak., 3861 
P. doniana J. Sm. ex Hook., 3934 

Aspidiaceae 

Ctenitis lanuginosa (Willd. ex Kaulf.) Copel., 3862 

Didymochlaena truncu/ata (Sw.) J. Sm., 3918 

Dryopteris inaequalis (Schlechtend.) Kuntze var. inaequalis, 3802, 3881 

D. kilemensis (Kuhn) Kuntze, 3880 

Tectaria gemmifera (Fe"e) Alston, 3685, 3895 

Polystichum zambesiacum Schelpe, 3801, 3925 

Aspleniaceae 

Asplenium aethiopicum (Burm. f.) Becherer, 3923 

A. blastophorum Hieron., 3688 

A. boltonii Hook, ex Schelpe, 3878 

A. christii Hieron., 3676 

A. dregeanum Kunze, 3856 

A. formosum Willd., 3696 

A. hypome/as Kuhn, 3885 

A. inaequi/aterale Willd., 3697 

A. monanthes L, 3855, 3858 

A. obscurum Bl., 3955 

A. rut i folium (Berg.) Kunze, 3854 



106 FERN GAZETTE: VOLUME 13 PART 2 (1986) 



A. unilaterale Lam., 3686 
A. ?volkensii Hieron., 3883 

A. sp. aff. gilpinae Bak., 3919 

Athyriaceae 

Athyrium scandicinum (Willd.) C. Presl. 3864 
Diplazium nemorale (Bak.) Schelpe, 3863, 3917 
D. pseudoporrectum Hieron., 3857 

Blechnaceae 

Blechnum attenuatum (Sw.) Mett, 3891 

B. ivohibense C. Chr., 3884 

B. punctu/atum Sw., 3813 

Cyatheaceae 

Cyathea humilis Hieron., 3886 

C. mossambicensis Bak., 3889 

Davalliaceae 

Nephrolepis biserrata (Sw.) Schott, 3974 

Dennstaedtiaceae 

Blot i el I a natalensis (Hook.) Tryon, 3675 

Histiopteris incisa (Thunb.) J. Sm., 3859 

Lonchitis occidentalis Bak., 3683 

Microlepia fadenii Pic. Ser., 3869 

M. speluncae (L.) Moore, 3932 

Gleicheniaceae 

Dicranopteris linearis (Burm. f.) Underw., 3709 

Grammitidaceae 

Ctenopteris sp. aff. villosissima (Hook.) Harley, 3815 

Grammitis kyimbilensis (Brause) Copel., 381 7A 

G. nanodes (A. Peter) Ching, 3817 

Xiphopteris strangeana Pic. Ser., 3818, 3847A 

Hymenophyllaceae 

Hymenophyllum polyanthos Sw. var. kuhnii (C. Chr.) Schelpe, 3866 

H. sibthorpioides (Bory ex Willd.) Mett. ex Kuhn, 3865 

H. splendidum v.d. Bosch, 3867 

Trichomanes borbonicum v.d. Bosch, 3868 

T. cupressoides Desv., 3681 

T. giganteum Bory ex Willd., 3682 

Lomariopsidaceae 

Bolbitis acrostichoides (Swartz) Ching, 3972 

B. auriculata (Lam.) Alston, 3661 

B. gemmifera (Hieron.) C. Chr., 3742A 

B. sp. aff. acrostichoides (Swartz) Ching, 3658, 3659, 3973 

Elaphoglossum acrostichoides (Hook. & Grev.) Schelpe, 3821 

E. macropodium (F£e) Moore, 3814 

E. phanerophlebium C. Chr., 3849 

E. spathulatum (Bory) Moore, 3659A 

Lomariopsis warneckei (Hieron.) Alston, 3890 

Polypodiaceae 

Be/visia spicata (Lf.) Mirb., 381 4A 

P/eope/tis excavata (Bory ex Willd.) Sledge, 3816 

Pteridaceae 

Pteris buchananii Bak. ex Sim, 3887 

P. preussii Hieron., 3888 

P. pteridioides (Hook.) Ballard, 3860 

P. quadriaurita Retz. spp. friesii (Hieron.) Schelpe, 3689 



ECOLOGY OF PTERIDOPHYTES IN MWANIHANA, TANZANIA 107 



P. sp. aff. mildbraedii/atrovirens, 3695 
P. sp. aff. prolifera Hieron, 3742 

Schizaeaceae 

Schizaea dichotoma (L) Smith, 3699 

Thelypteridaceae 

Amphineuron opulentum (Kaulf.) Holtt., 3743 

Christella gueintziana (Mett.) Holtt., 3946 

C. hispidula (Decne) Holtt., 3971 

C. sp. aff. gueintziana (Mett.) Holtt., 3694 

Menisorus pauciflorus (Hook.) Alston, 3655 

Sphaerostephanos arbuscula (Willd.) Holtt. ssp. africanus Holtt., 3693 

Pneumatopteris usambarensis Holtt., 3920 

Vittariaceae 

Antrophyum mannianum Hook., 3945 

Lycopsida 

Lycopodiaceae 

L dacrydioides Bak., 3807 

L ophioglossoides Lam., 3765 

L phlegmaria L., 3654 

L verticillatum L.f., 3808 

ACKNOWLEDGEMENTS 
We gratefully acknowledge the assistance of Dr B.S. Parris of the Royal Botanic 
Gardens, Kew, in naming the fern collection. The Tanzanian National Scientific 
Research Council and Ministry of Natural Resources and Tourism very kindly gave us 
permission to work in the Mwanihana Forest Reserve. The National Geographic 
Society and World Wildlife Fund generously supported the field work. Langson 
Kusoma and Henry provided much needed assistance in the field. 

REFERENCES 
DHV. 1982. Water supply survey: Southern Morogoro Region. DHV Consulting Engineers. 
LOVETT, J.C. (in press). Endemism and affinities of the Tanzanian montane forest flora. Ann. 

Miss. Bot. Gard. 
PARRIS, B.S. 1 985. Ecological aspects of distribution and speciation in Old World tropical ferns. 

Proc. Roy. Soc. Edin. 86B: 341-346. 
POCS, T. 1 976. Vegetation mapping in the Uluguru mountains (Tanzania, East Africa). Boissiera 

24: 477-498. 
WHITE, F. 1983. The Vegetation of Africa. UNESCO. 



108 FERN GAZETTE: VOLUME 13 PART 2 (1986) 



REVIEW 

INDEX FILICUM - SUPPLEMENTUM QUINTUM PRO ANNIS 1961-1975 by 
P.M. Jarrett, with T.A. Bence, J. W. Grimes, B.S. Parr is andJ.L.M. Pinner. Pp. 
245 Clarendon Press, Oxford. 1985. ISBN 0-19-854579-7. Price £25.00. 

Fern taxonomists have been wellservedby the provision of Indices givingthedetails of 
the publication of names. Index Filicum (1905-1906) was compiled by Carl 
Christensen who updated it by producing Supplements in 1913, 1917 and 1 934. This 
valuable work was continued by a committee of I.A.P.T. who published in 1 965 the 
Fourth Supplement covering the period from 1 934 to 1 960. 

The present Fifth Supplement has been compiled at Kew under the direction of Dr 
Frances Jarrett who undertook the onerous task of extracting names from the 
literature from 1 961 to 1 970, this being continued by her collaborators to completethe 
period covered by this work. 

The Supplement lists new names at all ranks between family and species but does 
not make taxonomic judgements as did Christensen's work. Infra-specific categories 
are not included as a general rule although they are quoted where they form the 
basionym for a new name. Where hybrids have been named their parentage is given 
and cross-referenced, for example Dryopteris x gotenbaensis Nakaike is quoted in full 
together with the information that it is the hybrid of D. hondoensis x uniformis. A 
further entry for D. hondoensis x uniformis refers the reader to the name D. x 
gotenbaensis. Thus, having either the name of the hybrid or its parentage the other can 
be found — a very useful feature when it increasingly is becoming common practice to 
give formal names to hybrids. 

All previous parts of the Index have dealt exclusively with ferns, a somewhat 
restrictive practice as fern workers are usually involved with the pteridophytes as a 
whole. Compilations of names for the fern allies have appeared in a variety of places at 
different times and it is a great convenience for them to be included along with the 
ferns as has been done in this Fifth Supplement for the first time. 

Users of the Index owe a great debt to the compilers for the meticulous way in 
which the work has been carried out. The publishers are also to be congratulated on 
the overall presentation and the quality of the type face. 

T.G. WALKER 



FERN GAZ. 13(2) 1986 109 



GEORGE GARDNERS PTERIDOPHYTE HERBARIUM 
AND LECTOTYPES OF GARDNER'S NEW FERN SPECIES 



B.S. PARRIS 
Botany School, Downing Street, Cambridge CB2 3EA, England* 

ABSTRACT 

George Gardner's original pteridophyte collections are at CGE and not with his 
phanerogam material (Brazilian specimens at BM, Ceylon specimens at K). 

INTRODUCTION 
George Gardner (1812-1849) is well-known as a botanical explorer in Brazil who later 
became superintendent of the Peradeniya Botanic Garden in Ceylon. His collections 
from both countries are large and important. The originals of his Brazilian material are 
stated to be in the British Museum (Natural History) (BM), while those of his Ceylon 
plants are said to be in the herbarium of the Royal Botanic Gardens, Kew (K) (Stafleu & 
Cowan 1976). These were presumably part of the herbarium offered for sale by 
Samuel Stevens and described in the Gardeners' Chronicle of 31 st May 1 851 (Anon. 
1 851 ). The disposition of Gardner's higher plants is undoubtedly as given above, but it 
is not so for the pteridophytes. Although both BM and K hold good sets of his 
pteridophyte numbers they are all duplicates and the original collections are held in 
the herbarium of the Botany School, University of Cambridge (CGE). 

A clue to how this may have come about is provided by a small printed label 
attached to some of the sheets which reads "Presented by Dr Churchill Babington". 
Dr Babington is known to have had an interest in cryptogams and presumably 
purchased the pteridophyte part of Gardner's collections either directly or indirectly 
from Samuel Stevens for his own use. In 1 865 he became Professor of Archaeology at 
the University of Cambridge At that time his cousin Charles Cardale Babington was 
Professor of Botany at Cambridge (Desmond 1977)and was acquiring duplicates of the 
major plant collections then available for the University Herbarium; doubtless 
Churchill Babington presented them to the Botany School as a gift to the herbarium 
which was rapidly expanding under his cousin's administration. 

The c. 1 990 sheets of Gardner's material are mounted on a distinctive blue lined 
paper and include not only his own gatherings but also duplicates from a variety of 
other collectors. I propose to document the latter in a subsequent paper. His own 
numbered original collections are represented, together with many un-numbered 
sheets which were presumably unicates or plants collected in too small a quantity to 
be numbered and distributed as duplicates. These are from Brazil, British Isles, Ceylon, 
India and Mauritius. None of the sheets of his own collections bear his name, but the 
handwriting is undoubtedly Gardner's and matches exactly that on his duplicates and 
in his notebooks kept at CGE, and the names and numbers on the duplicates (of which 
there are four incomplete sets in CGE) always correspond with those of the originals. 
The duplicates are widespread in herbaria (Stafleu & Cowan 1976) and usually have 
the plant name and number written in Gardner's hand; sometimes locality and date of 
collection is indicated, but ecological information is lacking. This is often present on 
the originals and is usually quoted verbatim in his descriptions of new species. 

Gardner described 1 6 new species of fern and now that his original pteridophyte 
collections have come to light (with the exception of one type) it seems expedient to 
choose lectotypes for them. 

*Present address: Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AE, England 



1 10 FERN GAZETTE: VOLUME 1 3 PART 2 (1 986) 



The lectotypes of George Gardner's fern species at CGE. 

Information in quotation marks is written in Gardner's hand on the herbarium sheets. 

Acrostichum alpestre Gardner 

in Fielding & Gardner, Sert.PI. t.25 (1844). 

[Gardner] "5924. On shady rocks near the summit of the Organ Mountains. March 1841. 

Acrostichum alpestre. Gardn." 

Lectotype chosen here. 

Adiantum filiforme Gardner 

in Hooker, Ic.PI. t.503 (1843). 

[Gardner] "2391. Shady cliffs of sandstone rocks. Oeiras, Piauhy. 1839. 

Adiantum filiforme, Gardn." 

Lectotype and 3 isolectotypes chosen here. 

Adiantum sinuosum Gardner 

in Hooker, Ic.PI. t.504 (1843). 

[Gardner] "3552. Serra de Natividade. Goyaz, Brazil. Jany. 1839. 

Adiantum sinuosum, Gardn." 

Lectotype and 3 isolectotypes chosen here. 

Anemia dentata Gardner 

in Fielding & Gardner, Sert.PI. ad t.70 (1844). 

[Gardner] "2387. Between Canabrava & Tranquiera, Province of Piauhy, Brazil. 1839. 

Anemia dentata, Gardn." 

Lectotype and 3 isolectotypes chosen here. 

Anemia glareosa Gardner 

in Fielding & Gardner, Sert.PI. t.70 (1844). 

[Gardner] "4086. Dry open campos, Goyaz, Brazil. Near Natividade and Arrayas. 1840. 

Anemia glareosa, Gardn." 

Lectotype and 2 isolectotoypes chosen here. 

Anemia pallida Gardner 

in Fielding & Gardner, Sert.PI. ad t.70 (1844). 

[Gardner] "3560 bis. On rocks in woods. Natividade, Goyaz. Jany. 1840. 

Anemia pallida, Gardn." 

Lectotype and 2 isolectotypes chosen here. 

Anemia wightiana Gardner 

in Calcutta Jour.Nat.Hist. 7:10, t.1-2 (1847). 

[Gardner s.n.] 

"Open rocky places on the Malabar slopes of the Neelgherries. Feby. 1845. 

Anemia wightiana. Gardner in Calcutta Journ." 

Lectotype and isolectotype chosen here. 

Asplenium woodwardioideum Gardner 
in Lond.Jour.Bot.(Hooker) 1 .547 (1842). 
[Gardner] "43. Corcovado, Rio de Janeiro. 1836. 
Asplenium woodwardioideum, Gardn." 
Lectotype chosen here. 

Cassebeera gleichenioides Gardner 

in Hooker, Ic.PI. t.507 (1843). 

[Gardner] "5295. Bushy rocky places. Diamond district. Aug. 1840. 

Cassebeera gleichenioides, Gardn." 

Lectotype and 2 isolectotypes chosen here. 

Cheilanthes monticola Gardner 

in Hooker, Ic.PI. t. 487 (1842). 

[Gardner] "3557. Summit of the Serra de Natividade. Province of Goyaz. Brazil. Jany. 1840. 

Cheilanthes monticola, Gardn." 

Lectotype and 3 isolectotypes chosen here. 



GEORGE GARDNER'S PTERIDOPHYTE HERBARIUM 1 1 1 



Coptophyllum buniifolium Gardner 

in Lond.Jour.Bot.(Hooker) 7. 133 (Jan. 1842). 

[Gardner] "4084. Serra de Natividade, Goyaz, Brazil, 1840. 

Coptophyllum buniifolium, Gardn." 

Lectotype and 3 isolectotypes chosen here. 

Coptophyllum millefolium Gardner 

in Lond.Jour.Bot.(Hooker) 1: 133 (Jan. 1842). 

[Gardner] "4083. Villa de Arrayas, Goyaz, Brazil, 1840. 

Coptophyllum millefolium, Gardn." 

Lectotype and 4 isolectotypes chosen here. 

Grammitis organensis Gardner 

in Hooker, Ic.PI. t.509 (1843). 

[Gardner] "5913. On rocks and on the stems of trees near the summit of the Organ Mountains, 

March. 1841. 

Grammitis organensis, Gardn." 

Lectotype and 2 isolectotypes chosen here. 

Polystichum pallidum Gardner 

in Lond.Jour.Bot.(Hooker): 7:547 (1842). 

[Gardner] "54. Woods, Corcovado, Rio de Janeiro, 1836. 

Polystichum pallidum, Gardn." 

Lectotype chosen here. 

Trochopteris elegans Gardner 

in Lond.Jour.Bot.(Hooker) 7/74 (Jan. 1842). 

[Gardner] "4085. Serra de Natividade, Goyaz, Brazil. Feby. 1840. 

Trochopteris elegans, Gardn." 

Lectotype and 3 isolectotypes chosen here. 

Only one species described by Gardner is not lectotypified here. Adiantum 
calcare urn Gardner in Hooker, Ic.PI. t. 467 (1 842) was based on Gardner no. 3551, but 
there is no original material of this number at present in CGE. Although duplicates 
exist (two of them in CGE) it is possible that the original may yet be found at CGE and 
typification should ideally wait until then. 

ACKNOWLEDGEMENTS 
I am grateful to Peter Sell, Assistant Curator of the herbarium, Botany School, 
Cambridge, for his advice and encouragement during my studies on the pteridophyte 
collections in CGE. 

REFERENCES 
ANON. 1851. Miscellaneous section. The Gardnerian Herbarium. Gardeners' Chronicle 1851: 

343-344. 
DESMOND, R. 1977. Dictionary of British and Irish Botanists and Horticu/tura/ists. Taylor & 

Francis: London. 
STAFLEU, F.A. & COWAN, R.S. 1976. Taxonomic Literature (2nd ed.) Volume 1: A-G. Bohn, 

Scheltema & Holkema: Utrecht. 



112 FERN GAZETTE: VOLUME 13 PART 2 (1986) 



REVIEW 

ILLUSTRIERTE FLORA VON MITTELEUROPA/GUSTAV HEGI Bd I Teil I 
PTERIDOPHYTA, 3rd fully revised edition by Karl U. Kramer. 309 pp., 1 1 
coloured plates, 275 figs. 200 x268mm. Paul Par ey, Berlin & Hamburg. ISBN 
3-489-500020-2. Price DM 228. (1984). (Pub/. 1983). 

Our concepts in species delimitations, distribution and general systematic relation- 
ships in Pteridophyta have changed considerably over the last 35 years and so this 
completely rewritten work (by J. Dostal, T. Reichstein, C.R. Fraser-Jenkins & K.U. 
Kramer) of this classic regional Flora, previously produced in 1935, is long overdue and 
greatly welcomed. Geographically the area of the Flora (shown in map-form onthe end 
papers) is difficult to equate with present-day political bounderies, being originally 
based on the German and Austrian Empires with the addition of Switzerland. Thus 
Alsace-Lorraine (France), Slovenija (Yugoslavia), Bohemia/Moravia (Czechoslovakia), 
Posen and Prussia (Poland) are included. This is hardly a floristic zone and one could 
ask why not include the whole of Poland and Czechoslovakia — and also Hungary? 

After a brief but concise general account of the group the book describes 39 
genera in 23 families. Comprehensive and detailed descriptions of genera and species 
are illustrated with line drawings and/or silhouettes and photographs of the plants in 
situ. Most species are shown in the eleven excellent coloured plates. Atthetaxonomic 
level, full synonymy, with references, and infra-specific variation is given; ecology, 
with associated species and both general (with maps) and detailed (in the area of the 
Flora) distribution is discussed. Vernacular names, including English, are also listed as 
are many relevant references to all chapters and subheads. Hybrids are treated less 
fully but still in considerable detail. 

A comprehensive work such as this, written by experts and edited with such 
thoroughness by Karl Kramer is, of course, a mile-stone in regional Floras. It will also 
be used as a reference book in a wider context e.g. for guidance to family names and 
classification. At this level it would have been useful to give synonymy of family names 
and a little discussion. In the reviewer's opinion the names and concepts used are 
sound and I hope that they will be taken up by other forthcoming European Floras. 

One cannot fault a book like this except on minor points. The illustrations vary: 
some are precise and pertinent e.g. in Polystichum; others not so helpful, e.g. the line 
drawings of Diphasiastrum do not show as much as the photographs and could have 
shown the important differences in lower leaves. Professor Kramer's eagle eye has 
eliminated even small errors but I noticed, under Ophioglossum azoricum, 0. 
vulgatum subsp. ambiguum (Cross, et Germ.) E.F. Warburg being quoted as published 
in C.T.W. Fl. Brit. Isles (1 962). Warburg published the combination in Watsonia 4: 41 
(1957). Much more unfortunate is the use of often unclear light microscope 
photographs of spores when surely SEM pictures would have said everything, and on 
Plate 5 the individual figures are incorrectly labelled in relation to the key on p. 1 60. 

A.C. JERMY 



FERN GAZ. 13(2) 1986 113 



A NEW DRYOPTERIS HYBRID FROM SPAIN 



CHRISTOPHER R. FRASER-JENKINS and MARY GIBBY 
Department of Botany, British Museum (Natural History), Cromwell Road, 

London SW7 5BD 

ABSTRACT 

A new Dryopteris hybrid, D. x asturiensis Fraser-Jenkins & Gibby, is described, and 
the cytology of this hybrid and D. corleyi Fraser-Jenkins is discussed. 

The northern coastal region of Spain is particularly rich in Dryopteris species, 
including the recently described endemic D. corleyi Fraser-Jenkins (1983) and D. 
guanchica Gibby & Jermy, known only from the Iberian peninsula and the Canary 
Islands, and thus provides opportunities for hybridization. A recently discovered hybrid 
from this area is D. x fraser-jenkinsii Gibby & Wid^n (1983), the hybrid between D. 
affinis (Lowe) Fraser-Jenkins subsp. affinis and probably D. guanchica, although the 
second parent could possibly beD. dilatata (Hoffm.) Gray. A second hybrid involving^. 
affinis has been found in N Spain recently, and although similar in morphology to D. x 
fraser-jenkinsii, differs in certain characters, and particularly in its cytology. Both 
hybrids are tetraploid, and produce some 8-celled sporangia with 164 bivalents at 
meiosis, and are capable of limited reproduction by spores, this apomictic character 
being inherited from D. affinis. However, D. x fraser-jenkinsii shows no chromosome 
pairing at meiosis in 16-celled sporangia, whereas the new hybrid has 41 bivalents 
and 82 univalents at first metaphase in 16-celled sporangia (Fig. 1). Morphological 
comparison suggests that the new hybrid may be D. affinis subsp. affinis x D. corleyi, 
with which it grows. D. corleyi is a tetraploid species (Fig. 2) that may have originated 
from the diploid species D. oreades Fomin and D. aemula (Ait.) 0. Ktze. (this is at 
present under investigation). Such a parentage could be compatible with the pairing 
seen in the hybrid, the bivalents being formed between the two 'oreades' genomes, 
one from D. affinis subsp. affinis and one from D. corleyi. 

Dryopteris x asturiensis Fraser-Jenkins & Gibby hybr. nov. 

(=D. affinis subsp. affinis x D. corleyi) 

Planta hybrida morphologia intermedia inter parentes. Stipes et rhachis crassiores 
quam in D. corleyi paleis fulvis basibus valde densioribus vestitis. Frons deltato- 
lanceolata, pinnata; pinnae sessiles, infimae e pinnulis basalibus longis gradatim 
decrescentes; pinnulae infernae valde sed non profunde lobatae usque ad dimidium 
latitudinis inter costam et marginem, lobi lati rectangulares, superi non-lobati vel lobis 
minoribus rotundatioribus; apices pinnularum late rotundati vel in plantis minoribus 
rotundato-truncati, lobi et apices pinnularum aliquot crenasvel dentes non manifestos 
ferentes, velut saepe apicibus acutis. Indusia valde decurvata ad margines eorum, 
aliquantum crassae et persistentia. Sporae abortivae, sed aliquot sporae magnae non- 
abortivae praesentiae. 

Holotype: Dryopteris x asturiensis Fraser-Jenkins & Gibby. c. 100m, in wood on 
sandstone 2km below Puron, c. 7km SE of Llanes, Oviedo to Santander, Oviedo 
(Asturias), Spain. C.R. Fraser-Jenkins 10835, 5 Oct. 1981 (BM) Fig. 3. 

Isotypes: Ditto (MA; Herb. T. Reichstein, Basel). 

Paratypes: c. 50m, in wood on sandstone above main Oviedo to Santander road, above 
Pendueles village, c. 2km E of Vidiago, E of Llanes, Oviedo to Santander, Oviedo 
(Asturias), Spain. C.R. Fraser-Jenkins 10778 (BM! K! MA! P!), 10779 (BM!), 10780 
(BM!), 10781 (BM!), 10798 (BM! Herb. M. Lainz, Gijdn University!). 

Plants occurring with both parents and also D. dilatata, D. affinis subsp. borreri 
(Newm.) Fraser-Jenkins and D. filix-mas (L.) Schott. Morphology intermediate 



114 



FERN GAZETTE: VOLUME 13 PART 2 (1986) 




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FIGURE 3 Dryopteris x asturiensis CRFJ 10835 holotype. 



A NEW DRYOPTERIS HYBRID FROM SPAIN 115 





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FIGURE 1a Dryopteris x asturiensis CRFJ 10835 spore mother cell from 16-celled sporangium 
at diakinesis showing 41 bivalents and 82 univalents; b explanatory diagram with bivalents in 
black, univalents outlined, x 750. 

FIGURE 2a Dryopteris corleyi CRFJ 10782 spore mother cell at diakinesis showing 82 bivalents; 
b explanatory diagram, x 750. 

between the parents. Stipe and rhachis thicker than in D. corleyi and markedly more 
densely clothed in light brown scales with dark bases. Frond deltate-lanceolate, once 
pinnate; pinnae sessile, the lowest ones gradually tapering from long basal pinnules; 
lower pinnules markedly but shallowly lobed up to half the width between the midrib 
and margin with wide, rectangular lobes, upper pinnules unlobed or with smaller, 
more rounded lobes; pinnule apices broadly rounded, or in smaller plants, rounded- 
truncate, the lobes and pinnule apices bearing a few insignificant cren at ions or obtuse 
teeth, though often with pointed apices. Indusia strongly curved down at their 
margins, somewhat thick and persistent. Spores abortive but with a few large, good 
spores present. Differs from D. affinis in its long stipe, deltate-lanceolate frond, long 
lowest opposite pair of pinnules on the lower few pairs of pinnae, much more deeply 
lobed pinnules, the lower ones with narrower apices, and mostly abortive spores. 
Differs from D. corleyi in its denser and browner scales, many with dark bases, its 
thicker stipe and rhachis, more sessile pinnae and pinnules, less lobed lobes or divided 
pinnules, markedly broader pinnule apices, thicker indusia and mostly abortive spores. 
Differs from D. xfraser-jenkinsii in its slightly paler stipe scales, markedly broader, less 
pointed pinnule apices, more rounded pinnule lobes with shorter, less acuteteeth, and 
a generally flatter lamina with less twisted segments. 



•, -, 6 FERN GAZETTE: VOLUME 1 3 PART 2 (1 986) 



REFERENCES 
FRASER JENKINS, C.R. 1982. Dryopteris in Spain, Portugal and Macaronesia. Bol. Soc. Brot, 

Ser. 2a. 55: 175-336. 
GIBBY, M. & WIDEN, C.-J. 1983. Three new Dryopteris hybrids from Spain and the Canary 
' Islands. Fern Gaz. 12: 267-270. 



REVIEWS 

MED-CHECKLIST 1 PTERIDOPHYTA (ed. 2) GYMNOSPERMAE 
DICOTYLEDONES (ACANTHACEAE-CNEORACEAE) edited by W. Greuter, 
H.M. Burdet & G. Long. 330+ C pp. 312 x210mm. Published by Conservatoire 
et Jar din botaniques, Ville de Gendve, Med-Checklist Trust of OPTIMA, 
Gendve ISBN 2-8277-0151-0, ISBN 2-8279-0004-1 , 1984. 

This is a checklist of vascular plants from all the countries that border on the 
Mediterranean, plus Portugal, Bulgaria, the Crimea and Jordan. The taxonomic 
advisers for pteridophytes are C.R. Fraser-Jenkins, A.C. Jermy and T. Reichstein, and 
the book includes over 150 fern species. For each species the name, authority and 
source is given and synonyms, and the distribution (by country) within the 
Mediterranean region. For recent or doubtful records the reference is given in the 
appendix. Unfortunately some of the names used in this checklist are not those found 
in Hegi's lllustrierte Flora von Mitteleuropa (Kramer 1983; see review p. 1 1 2) which 
was published in the same year, and this must reflect an editorial decision, since C.R. 
Fraser-Jenkins and T. Reichstein were pteridophyte advisers for Hegi. For example, 
the Med-Checklist uses Lycopodium for all the Lycopodiaceae, whereas Hegi uses 
Lycopodiella, Diphasiastrum etc., and such disagreement is frustrating for all 
pteridologists. However, the Med-Checklist, which uses a computerized system for 
information processing, formating etc. and thus minimizing typographical errors, 
provides a valuable reference source. 

M. GIBBY 



EVOLUTIONARY CLAD/ST/CS OF MARA TT/ALEAN FERNS by Christopher R. 
Hill & Josephine M. Camus. Bulletin of the British Museum (Natural History), 
Botany series Vol. 14 No. 4. 27 February 1986. Price £14.50. 

The application of cladistic methods to problems of fern taxonomy has not, as yet, been 
widespread. It is encouraging, therefore, to read this paper on the classification of 
Marattiales which relies on cladistics to resolve relationships within the order. The 
concepts and terminology of the method are given, which is useful for those readers 
unfamiliar with cladistics. All species of the order have not been included, but the 23 
species selected cover the range of variation in each of the extant genera; 73 
characters are used ranging from the stelar anatomy of the adult stem to the shape of 
the exine spines of the spores. The resulting cladogram indicates the distinctness of 
both Christensenia and Danaea, and the paraphyletic nature of Marattia, 
Macroglossum, Angiopteris and Archangiopteris. As a check on the taxonomic 
relationships defined by the cladogram the authors have used stratigraphy, phyto- 
geography and ontogeny, and present a convincing case for their conclusions. I look 
forward very much to seeing their further work on revising the genera and species of 
Marattiaceae. 

B.S. PARRIS 



FERN GAZ. 13(2) 1986 117 



SUBGENERIC NAMES IN SELAGINELLA 



A. C. JERMY 
British Museum (Natural History), Cromwell Road, London SW7 5BD 

ABSTRACT 

In an account of Selaginellaceae to be published in The Families and Genera of 
Plants: Pteridophyta (ed. Kramer et al.) the author proposes five subgenera in the 
sole genus Selaginella: Selaginella, Ericetorum Jermy, Tetragonostachys Jermy, 
Stachygynandrum (P. Beauv.) Baker and Heterostachys Baker. In this brief synopsis 
the names Ericetorum Jermy and Tetragonostachys Jermy are described and 
validated. 

INTRODUCTION 
The genus Selaginella contains approximately 700 species for the most part 
concentrated in the tropical areas of the world. Botanists of the late eighteenth century 
(e.g. Adanson, Boehmer, Palisotde Beauvoir)describing the relatively small number of 
species available to them at that time saw generic distinctions that cannot be upheld 
today. Kuntze (1891), in the belief that Lycopodioides Boehmer was the earliest name 
forthe genus, published some 320 newcombinations inthat genus. Rothmaler (1 944) 
proposed acceptance of these earlier generic names but large-scale nomenclatural 
changes have not been published without the broad revision of the classification of the 
family that is needed. The present author, having studied Selaginella in some detail 
throughout its range, has proposed in a forthcoming book (The Families and Genera of 
Plants: Pteridophyta, ed. K.U. Kramer et al.) the following infrageneric classification 
which is published here in synopsis only in order to validate two of the names used. A 
full account of the morphology and relationships of these taxa is in preparation. 

SELAGINELLA P. Beauv. 
Palisotde Beauvoir, Magasin Encyc!. 5:478(1804); Prod. fam. Aetheog. 101 (1805), 
em. Spring in Flora (Regensb.) 21: 148 (1838); nomen conserv. 

Subgenus Selaginella. Type-species Selaginella spinosa P. Beauv. = Selaginella 
selaginoides (L.) Link. 

Syn.: Subgen. Homoeophyllum (Spring) Hieron. & Sadeb. in Engler & Prantl, Nat. 
Pflanz. 1 (4): 669 (1902) p.p. 

Stems erect, new primary shoots arising from the base upon maturation of the single 
terminal strobilus, rooting from a basal hypocotular node; leaves and sporophylls 
spirally arranged, uniform and herbaceous. 

Two species: S. selaginoides — circumboreal in the Northern Hemisphere, south to the 
Canary Islands; S. deflexa Brackenridge — endemic to Hawaiian Islands. 

Subgenus Ericetorum Jermy subgen. nov. 

Syn.: Subgen. Homoeophyllum (Spring) Hieron. & Sadeb. in Engler & Prantl, Nat. 
Pflanz. 1 (4): 669 (1902) p.p. 

Rami erecti, aut ramosi aut rarius simplices, e caule repenti solenostelam continent! 
procumbente, exorientes; folia aequabilia, saltern ad basin decussata, plus minusve 
herbacea, lamina ovata vel ovato-lanceolata; sporophylla tetrasticha. 
Type-species Selaginella uliginosa (Labill.) Spring, Bull. Acad. R. Belg. 10: 136(1843). 
Stems erect, either unbranched or more compound, arising from a creeping 
solenostelic stem; leaves uniform, decussately arranged at least below, more or less 
herbaceous with an ovate or ovate-lanceolate lamina; sporophylls tetrastichous. 
Three species: S. uliginosa — Australia and Tasmania; S. gracillima (Kunze) Spring — 
SE Australia; S. pygmaea (Kaulf ) Alston — southern Africa. 



118 FERN GAZETTE: VOLUME 13 PART 2 (1986) 

Subgenus Tetragonostachys Jermy subgen. nov. 

Syn.: Subgen. Homoeophyllum (Spring) Hieron. & Sadeb. in Engler & Prantl, Nat. 

Pflanz. 1 (4): 669 (1902) p.p. 

Planta repens, caulibus pleuriramosis, per totam longitudinem radices emittentibus, 
prostrata et saepe tegetes formans, vel humilis ramis erectis; folia spiraliter disposita, 
aequabilia vel in ramis prostratis dimorphescentia, plerumque coriacea, lineari- 
lanceolata, apice aliquando acicularia vel apicepilum praedita;sporophyllatetrasticha. 
Type-species Selaginella rupestris (L) Spring, Flora 21: 149 and 182 (1838). 
Plants creeping, stems much-branched, rooting throughout their length, prostrate and 
often mat-forming or with short erect branches; leaves spirally arranged, similar, or on 
prostrate branches slightly dimorphic, usually coriaceous, linear-lanceolate, apex 
sharp or bearing a hair; sporophylls tetrastichous. 

About 50 species ranging from southern North America through the tropics of South 
America, Africa and the Indian subcontinent to N China and Japan. 

Subgenus Stachygynandrum (P. Beauv.) Baker, J. Bot., Lond. 21:3 (1883) emend. 
Jermy, Fern Gaz. 13:1 18 (1986). Basionym: Stachygynandrum P. Beauv. ex Mirbel in 
Lam. & Mirbel, Hist. Nat. Veg. 3: 477 (1802). Type-species Lycopodiumflabellatum L.= 
Selaginella flabellata (L.) Spring. 

Syn.: Subgen. Heterophyllum Hieron. & Sadeb. in Engler & Prantl, Nat. Pflanz. 1 (4): 
673 (1902); subgen. Homostachys Baker, J. Bot., Lond. 21: 4 (1883). 

Stems either creeping with prostrate branches, or erect with various and often complex 
branching systems, leaves dimorphic, at least on the secondary branches, in four 
distinct rows, those of the upper rows being distinctly smaller; sporophylls uniform, or 
in a few cases showing slight dimorphism, tetrastichous. 

Baker's concept is enlarged to include those species (S. ciliaris (Retz.) Spring; S. 
pallidissima Spring) he separated as being in his fourth subgen., Homostachys. Those 
species and others, such as those included in that subgenus by Walton & Alston (1938), 
have loose strobili with sporophylls that begin to show some dimorphism. In my 
opinion, and in that of N. Quansah (pers. com m.) who has studied the African species in 
detail these can rightly be included in subgen. Stachygynandrum. 
About 600 species ranging throughout the tropics of all continents. 

Subgenus Heterostachys Baker, J. Bot., Lond. 21: 4 (1883). Lectotype-species 
Selaginella heterostachys Baker, J. Bot., Lond. 23:177 (1885). 

Syn. subgen. Heterophyllum Hieron. & Sadeb. in Engler & Prantl, Nat. Pflanz. 1 (4): 673 
(1902). 

Stems creeping and much-branched, or secondary branches erect and suffruticose, 
rooting in branch axils; leaves as in subgen. Stachygynandrum; strobili complanate, 
sporophylls dimorphic, tetrastichous, those on the ventral side smaller than those on 
the upper side of the shoot. 
About 60 species with a distribution range similar to that of subgen. Stachygynandrum. 

REFERENCES 
KUNTZE, 0. 1891. Revisio Generum Plantarum, 1 : 824-827. 

PALISOT DE BEAUVOIR. 1805. Prodromus de la famille Aetheogamie. Pp. 112. Paris. 
ROTHMALER, W. 1944. Pteridophyten-Studien I. Feddes Repert. 54: 55-82. 
WALTON, J. & ALSTON, A.H.G. 1938. Lycopodiinae. Chap. XVII in Manual of Pteridology (ed. F. 
Verdoorn). Nijhoff, The Hague. 



FERN GAZ. 13(2) 1986 119 

SHORT NOTES 

THE OCCURRENCE OF SCHIZAEA DICHOTOMA IN TANZANIA 

The curious and primitive fern Schizaea dichotoma (L.) Smith has been discovered 
recently in the Uzungwa mountains of Tanzania in the Mwanihana Forest Reserve 
above Sanje village at latitude 7°50'S and longitude 36°55'E. It is now known from 
three collections, which represent the first records of the species in continental Africa. 
Its previous known rangewas Madagascar to Polynesia and Australia (Holttum, 1 968), 
where it is widespread and frequently collected. 

In Tanzania it grows on the forest floor in leaf litter under a canopy of Albizia 
gummifera, Filicum decipiens, Funtumia africana, and Parinari excelsa, with the 
saprophytic herb Seychellaria africana, at an altitude of between 900-1 200m. The 
forests themselves occur on the east facing slopes of the ancient crystalline Uzungwa 
mountains in an annual rainfall of 2000-2500mm, and are well known for the high 
degree of endemism they contain (Lovett, in press). 

In view of the previous known range of Schizaea dichotoma, the question arises as 
to whether its distribution pattern is a relict which predates the break up of 
Gondwanaland and the separation of Madagascar from continental Africa, or whether 
its occurrence in Tanzania is a result of later long distance dispersal from Madagascar. 
The former idea has been suggested to explain the distribution of the bryophyte family 
Rutenbergiaceae which also occurs on the ancient crystalline East African mountains 
and Madagascar (Pocs, 1975). However, for Schizaea dichotoma, long distance 
dispersal is perhaps more likely in view of its limited distribution in Tanzania, and its 
spores might be carried by the prevailing winds from Madagascar to Tanzania. 

Interestingly enough the associate of Schizaea dichotoma, Seychellaria africana, 
which is currently only known from the Uzungwa mountains, belongs to a genus 
which is otherwise only found on Madagascar and adjacent islands (Vollesen, 1 980). 
Perhaps it too reached Tanzania by long distance dispersal of its small seeds. The 
association of Schizaea and Seychellaria is even more curious in that it has been noted 
in the Neotropics that Schizaea is found in association with saprophytic plants in the 
same family as Seychellaria, the Triuridaceae (Maas, 1979), and the fact that this 
condition is also found in Tanzania may suggest that Schizaea is also saprophytic to 
some degree. 

I am grateful for the assistance of Barbara Parris in the identification of Tanzanian 
ferns, and the Tanzania National Scientific Research Council who very kindly gave me 
permission to conduct field work in Tanzania. The National Geographic Society and 
World Wildlife Fund generously provided the support for the field work. 

Collection numbers from the Tanzanian locality are: Lovett 244, (K, DSM) Stuart & 
Rodgers 838 (K, DSM), Thomas 3699 (K, MO). 

REFERENCES 
HOLTTUM, R.E. 1968. Ferns of Malaya. Flora of Malaya Vol. II, Ed. 2. 
LOVETT, J.C. (In press). Endemism and affinities of the Tanzanian montae forest flora. Ann. Miss. 

Bot. Gard. 
MAAS, P.J.M. 1979. Neotropical saprophytes. In: Tropical Botany. Ed. Kai Larsen & Lauritz B. 

Holm-Nielsen. Academic Press. 
POCS, T. 1975. Affinities between the bryoflora of East Africa and Madagascar. Boissiera 24: 

125-128. 
VOLLESEN, K. 1 980. A new species of Seychellaria (Triuridaceae) from Tanzania. KewBull. 36: 

733-736. 

J.C. LOVETT 
Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166, U.S.A. 



1 20 FERN GAZETTE: VOLUME 1 3 PART 2 (1 986) 



A CHROMOSOME COUNT FOR ANOGRAMMA LEPTOPHYLLA 

IN MADEIRA 

A comprehensive study of the cytology of the native ferns of Madeira has recently been 
completed (Manton et al. 1986). However, for one fern, Anogramma leptophylla (L.) 
Link, a definitive count proved difficult, owing in part to a tendency for clumping of the 
chromosomes but also to the annual habit of the fern. In early Spring of this year J.F.M. 
and M.J. Cannon made fixings in the wild (Madeira: Levada from Bocodo Riscoeastto 
Canical tunnel entrance. Cannon & Cannon 5169, 10 March 1986. BM), and a 
chromosome count of 26 bivalents at meiosis has now been confirmed (Fig. 1 ). This is 
in agreement with Fabbri (1963) quoting Tutin as 'n= 26?' for material from Jersey, 
and Kurita (1 971 ) who gives n = 26 for material from Europe without giving a precise 
locality, and contrasts with counts of n = 29 for this species from New Zealand 
(Brownlie 1 958) and South Africa (Baroutsis & Gastony 1 978), and n=56-7 (Mehra & 
Verma 1960) and n=58 (Verma, quoted by Manton et al. 1986) from India. 






t 



4 



mm 



• 



FIGURE 1a Spore mother cell of Anogramma leptophylla at metaphase 1 showing 26 bivalents; 
b explanatory diagram, x 1000. 



REFERENCES 

BAROUTSIS, J.G. & GASTONY, C.J. 1978. Chromosome numbers in the genusAnogramma. II. 

Am. Fern J. 68: 3-6. 
BROWNLIE, G. 1958. Chromosome numbers in New Zealand ferns, frans. R. Soc. N.Z. 85: 213- 

216. 
FABBRI, F. 1963. Primo supplemento alle Tavole cromosomiche delle Pteridophyta di Alberto 

Chiarug i. Caryo/ogia, 16: 237-335. 
KURITA, S. 1971 . Chromosome study of four species of leptosporangiate ferns. Ann. Rep. Foreign 

Students' Coll., Chiba Univ. 6: 41-43. 
MANTON, I., LOVIS, J.D., VIDA, G. & GIBBY, M. 1 986. Cytology of the fern flora of Madeira. flu//. 

Br. Mus. nat. Hist. (Bot.) 15: in press. 
MEHRA, P.N. & VERMA, S.C. 1960. Cytological observations on some west Himalayan 

Pteridaceae. Caryo/ogia 13: 613-650. 

MARY GIBBY 

Department of Botany, British Museum (Natural History), 

Cromwell Road, London SW7 5BD 



FERN GAZ. 13(2) 1986 121 



CYSTOPTERIS IN THE CAPE VERDE AND CANARY ISLANDS 

The specimens of Cystopteris which I collected in the Cape Verde Islands have been 
published by me as C. fragilis sens. lat. (Lobin, 1982). 

Now, using the vein character of Rocha Afonsa (1 982) and the spore characters I 
myself have observed, I can say that the Fogo (Cape Verde Islands) specimens are C. 
fragilis (L.) Bernh. and that the specimens which I collected on Tenerife (Canary 
Islands) are C. viridula (Desv.) Desv. The key characters I used are: 

Veins ending in the apex of the teeth; spores echinate, loosely covered with 

spines ± equal in size and shape C. fragilis 

Veins ending in the sinus between two teeth; spores spiny-lacunar, so densely 
covered with spines unequal in size and shape and connected at their base, that 

they obscure the surface C. viridula 

C. fragilis was discovered in the Cape Verde Islands in 1863 by A. Stubel (Bolle, 
1866). Asecond collection of this rarefernwas madebyChevalier(1935)andthethird 
by Barbosa (Nogueira, 1 975). I found the species growing at Fogo in the walls of small 
shady humid valleys (ribeiras) at about 800to 1200m alt. and apart from one specimen 
in the southwest, all on the north side. 

REFERENCES 
BOLLE, C. 1 866. Die Standorte der Farm auf den Canarischen Inseln III. Z. ges. Erdk., 1: 209-238; 

Berlin. 
CHEVALIER, A. 1 935. Leslies du Cap Vert. Flore de I'Archipel. Rev. int. Bot. appl. Agric. trop., 15: 

733-1090: Paris. 
LOBIN W. 1982. Untersuchung u'ber Flora, Vegetation und biogeographische Beziehungen der 

Kapverdischen Inseln. Cour. Forsch.-lnst. Senckenberg, 53: 112 pp.; Frankfurt. 
NOGUEIRA, I. 1 975. Plantas colhidas pelo Eng.° L.A. Grandvaux Barbosa no arquip^lago de Cabo 

Verde I. Pteridophyta. Garcia Orta, (Bot.) 2 (2): 79-84; Lisbon. 
ROCHA AFONSA, M. 1982. Contribuic3o para o estudo do g^nero Cystopteris Bernh. em 
Portugal continental e insular. Bo/. Soc. Brot, (2) 55: 337-352; Coimbra. 

WOLFRAM LOBIN 
Forschungsinstitut Senckenberg, Senckenberganlage 25, 

D-600 Frankfurt, W. Germany 

CYSTOPTERIS VIRIDULA IN MAINLAND AFRICA 

Although known from the Cape Verdes, Macaronesia, south-west Europe, west 
Mediterranean Europe, Morocco and Algeria, Cystopteris viridula (Desv.) Desv. has not 
so far been reported from mainland Africa (in the floristic sense). My father and I 
recently found it on Mount Oku in North-West Province, Cameroon: above road on NE 
side of Mt. Oku, c. 4km SE of Oku on Kumbo road, NE of Bamenda, c. 2300m alt., on 
moss-covered boulder in dense natural forest by stream, above encroaching potato 
fields. Coll: CDFJ & CRFJ 1 1489, 6 June 1985 (BM, H). 

This suggests that this species could be yet another example of the African affinity 
of much of the Macronesian and Atlantic European floristic element in ferns, which I 
see as secondarily invading south-western Europe in many cases, though often with 
presumed more ancient Asian connections, sometimes via Africa. I have not yet been 
able to investigate other west or central African populations of the Cystopteris fragilis 
(L.) Bernh. group from the Tibesti, Hoggar etc. 

ACKNOWLEDGEMENTS 

The author would like to thank his father, CD. Fraser Jenkins of Bridgend, the 

Botanical Museum, University of Helsinki and Professor T. Reichstein for their 

generous financial help. c R FRAS ER-JENKINS 

c/o Department of Botany, British Museum (Natural History), 

Cromwell Road, London SW7 5BD 



1 22 FERN GAZETTE: VOLUME 1 3 PART 2 (1 986) 



FIRST RECORD OF EQUISETUM x DYCEI 
IN CONTINENTAL EUROPE 

During a botanical survey of plants growing in the "Maximilianpark" in the vicinity of 
Hamm (North Rhine-Westphalia, Federal Republic of Germany) one of us (U.P.) 
collected a strange looking horsetail which at first was thought to represent Equisetum 
x litorale Kuhlew. ex Rupr. Further studies revealed, however, that the plants showed 
closer relationships to E. palustre L. than to E. arvense L. Our tentative determination 
as£. x dycei C.N. Page was subsequently confirmed byC.N. Pagetowhom we sent our 
plants for examination. As far as we know this is the first record of £. x dycei outside the 
British Isles and the first for continental Europe. Diagnostic features of this hybrid are 
given by Page (1982). 

The environmental conditions of the E. x dycei stand in Hamm aresimilartothose 
described by Page (1985) for the British and Irish localities. It is a wet habitat being 
permanently flooded by shallow water and has been disturbed or at least modified in 
some degree by man. This occurred recently duringthe development of the whole area 
for the "Landesgartenschau", an open air display of ornamental flowers exhibiting 
modern trends and features of garden architecture, which took place in 1 984. In the 
course of the work the site where E. x dycei grows was covered with loamy soil and an 
artificial water regime installed to keep the site flooded. Additionally, several species 
of macrophytes (e.g. Lythrum sa/icaria, Typha angustifolia, Carex acutiformis) were 
planted to re-create a vegetation cover. Both parental species of E. x dycei, E. palustre 
and£. f/uviati/eL., are present, but only a small number of shoots; these have not been 
planted. The vegetation is still very scarce and open, and this is another common 
feature with the Scottish and Irish localities. Here the rather few (about 20), small and 
weakly growing shoots of E. x dycei persist. This hybrid appears to be a weak 
competitor and disappears as soon as other plants start to grow more vigorously and 
vegetation becomes more dense. This is very much in contrast to E. x litorale which can 
form large colonies and seems to compete quite successfully with many other plant 
species. The whole circumstances suggest that the E. x dycei stand in Hamm 
originated there only a few years ago. 

Although it is difficult to make any predictions on the future development of the 
population we are inclined to believe that the plants will not survive for a very long 
time. The stand could be adversely affected by both enhanced growth of other 
macrophytes and possible disturbance of the habitat by recreational activities. 

Regarding the distribution of E. x dycei we fully agree with the statement of Page 
(1 985) that E. x dycei can very easily be overlooked and it may well turn out that it is 
more frequent on the European continent than this one reported locality suggests. 



ACKNOWLEDGEMENTS 
We thank Dr C.N. Page, Edinburgh, for his kind support and for examining our 
Equisetum specimens and Mr Ralph David, Witten, for improving our English. 



REFERENCES 
PAGE, C.N. 1982. The ferns of Britain and Ireland. Cambridge University Press. 
PAGE, C.N. 1985. The distribution and habitats of Equisetum x dycei. Pteridologist 1 :60. 



H.W. BENNERT & U. PETERS 

Spezielle Botanik, Ruhr-Universitat Bochum, Universitatsstrasse 150, 

P.B. 10 21 48, D-4630 Bochum 1, West Germany 



FERN GAZ. 13(2) 1986 123 



AN ANOMALY OF BOTRYCHIUM MATRICARIIFOLIUM 

In August 1 985 the author found one specimen of Botrychium matricariifolium A.Br, 
ex Koch in the hills of the Ceskomoravska' Vrchovina about 70km northwest of Brno 
(Czechoslovakia). It was surprising to see that it bore sporangia on the sterile part of its 
frond. 

Since this specimen was rather old when it was found it was impossible to 
determine the correct species at once so it was sent to Prof. Dr. Follmann (Koln) for 
detailed analysis. He ascertained the abnormal sporangia to be stunted and the 
number of chromosomes to be about 180. As B. virginianum (L) Sw. must be 
eliminated because of its very different habit the specimen can only be B. 
matricariifolium. 

For further details the author may be contacted. Special thanks are due to Prof. Dr. 
Follmann for his efforts. 

HARALD SCHUMANN 
Bentelerstrasse 8, D-4400 Munster, West Germany 



CORRECTIONS TO INDEX FILICUM 

Index filicum is now produced by the Royal Botanic Gardens, Kew, and as the new 
editor I am keen to correct the errors in the original and subsequent supplements, lam 
aware of a number of these but am sure that others are as yet undetected. I very much 
welcome information on any omissions and mistakes known to my colleagues. 
Infraspecific names are to be included in the next supplement, which will run from 
1 976-1 985. From 1 986, new names of pteridophytes will be published annually, as an 
appendix of an annual Index Kewensis, but Index Filicum will continue to be published 
at 5 or 10 year intervals. 

B.S. PARRIS 
The Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AE 



124 FERN GAZETTE: VOLUME 13 PART 2 (1986) 

OBITUARY 

PROFESSOR E.A.C.LE. SCHELPE 

Professor Edmund Andre" Charles Lois Eloi (Ted') Schelpe died after a sudden heart 
attack at his home on Saturday 12th October 1985. Professor Schelpe was born in 
Durban on 27th July 1924. He completed a BSc with distinction in Botany at Natal 
University in 1 943 and at the same University obtained an MSc (Class I) in 1 946. He 
obtained the DPhil degree of Oxford University (Wadham College) in 1 952. He heldthe 
following posts at the University of Cape Town: Lecturer in Botany, 1953-1954; Senior 
Lecturer and Curator of the Bolus Herbarium, 1954-1958; Associate Professor and 
Curator of the Bolus Herbarium, 1968-1973, the title Curator being changed to 
Director from 1970; Professor (ad hominem) and Director of the Bolus Herbarium, 
1973-1985. Professor Schelpe was an elected Fellow of the Royal Society of South 
Africa, the Linnean Society of London, andthe University of Cape Town. From 1 964 he 
had been a Member of the Committee on Pteridophyta of the International Association 
of Plant Taxonomists. Professor Schelpe has made major contributions to the 
systematics of Pteridophyta in southern Africa. A few days before he passed away he 
finished checking the final proofs of the Pteridophyta volume for the Flora of Southern 
Africa. This was a culmination of studies in the course of which he published the 
accounts of the Pteridophyta for the Flora Zambesiaca, the Flora de Mccambique and 
the Conspectus Florae Angolensis. He published over 100 scientific papers and books, 
which besides Pteridophyta covered topics in orchids, bryophytes, plant ecology and 
phytogeography. Professor Schelpe was a distinguished and internationally well- 
known botanist and his passing away is a sad loss to the scientific community. 

Information supplied by A.V. HALL 
Bolus Herbarium, Rondebosch, 7700 South Africa 



FERN GAZ. 13(2) 1986 125 



REVIEWS 

FERNS OF JAMAICA by G.R. Proctor. 631 pp., 135 figures, 22 maps, 175 x 
250mm. British Museum (Natural History), London. ISBN: 0-565-00895- 1 
1985. Price £50.00. 

Jamaica is an island especially rich in ferns for which no adequate modern handbook 
was available. The defect has now been remedied in a very authoritative manner by the 
publication of this book. All groups of pteridophytes are covered and some idea of the 
richness of the flora and the techical problems facing the author can be gained from 
the fact that 579 species and a further 30 clear varieties are described. 

A very valuable feature of the short Introduction is the listing of the 1 78 collectors 
known to have gathered ferns in Jamaica, together with their collecting dates and 
herbaria where the author has seen their specimens. 

The main text is easy to read and well laid-out —thus for each family there is a 
general account and a note on any special literature, followed by clear keys to the 
genera. The species entries are concise and informative, giving the detailed authority, 
the type specimen, basionym and synonyms, species description, general range and 
Jamaican distribution and a short entry on habitat, together with comment as to 
relative abundance in Jamaica. 

The specific descriptions average some 10-12 lines and are clear proof of Dr 
Proctor's familiarity with the plants in the field. Use has been made of published 
cytological evidence wherethis hasthrown light on a particulartaxonomic aspect such 
as the separation of taxa and the recognition of hybrids, etc. 

In taking a wide view of such genera as Thelypteris, Polypodium, Grammitis, 
Cyathea, etc., and by maintaining a hierarchy based on the subgenus, Proctor is of the 
opinion that the ends of classification are thus best served. All the keys are clear and 
well-constructed and avoid the use of comparative terms which are an unfortunate 
feature of some works and are of little use unless the student has both taxa to examine. 
The acid test of any key is how easy and accurate it is in use and I selected eight of the 
largest genera and from these pulled out at random folders of 20 species, all of which 
keyed out satisfactorily. There was one discrepancy, namely in the key leading to 
Polypodium loriceum where the rhizome was said to be "bearing a few scattered 
appressed scales, otherwise naked" whilst the specific description (p. 529) describes 
the rhizome as bearing numerous appressed scales. Indeed, in the same folder of P. 
loriceum that I examined, some specimens had numerous rhizome scales whilst 
others had very few. Nevertheless despite this difficulty the keying process still gave 
the right answer because of the other alternatives. 

A delightful feature of the book is in the illustrations. Seventy five of the 83 native 
genera are delineated and a very skilful choice has been made to demonstrate a range 
of artistic styles and printing methods — some old such as those from Rovirosa, 
Hooker, Schkuhr, etc., and others modern from Small, Stolze and A.R. Smith for 
example. In a number of cases excellent original drawings by P.J. Edwards are 
included. 

In summary, this is an invaluable work written by the leading authority on 
Jamaican ferns who has built up an intimate knowledge of these plants in the field 
over many years and who has put this to use in a very clear and concise manner. No 
one who has an interest in these ferns can affordto bewithoutthis book, althoughthe 
price is somewhat daunting. I hope that we may look forward to sequels covering the 
history of plant collecting in Jamaica and an account of the ecology and behaviour of 
these ferns. 



12 6 FERN GAZETTE: VOLUME 13 PART 2 (1986) 



It may be somewhat churlish to adversely comment on a feature of such an 
excellent book, especially as it is not the responsibility of the author, but mention must 
be made of the binding Books of this type which are going to be used extensively both 
in the laboratory and the field will be subject to quite heavy wear and tear. A first 
essential is to provide covers which will stand up to this and which preferably should 
be waterproof or wipeable. In the short time I have had my copy (which has only been 
used in the laboratory) the cover is showing dirty marks and on attempting to wipe 
them off the red dye has come away. In addition the corners are fraying. Surely more 
thought could be put by publishers to the use to which a book is going to be put and 
provide more appropriate covers, particularly in view of the high price of books in 

general. 

T.G. WALKER 



GAMETOPHYTES OF OPHIOGLOSSACEAE by D.D. Pant, D.D. Nautiyal and 
D.R. Misra. Phyta Monograph 1:1-111. 1984. 180 x250mm. Published by the 
Society of Indian Plant Taxonomists, Allahabad, 211002. India. Price not 

given. 

This is a thorough descriptive account of the gametophytes of six species of 
Ophioglossum, four of Botrychium and Helminthostachys zeylanica, and an historical 
review of work on the subject. The gametophytes of Ophioglossaceae are geophilous, 
cylindrical, fleshy and tuberous, and for the most part monoecious. Xylem stands are 
shown to be present in the centre of the prothallus of H. zey/anicum and comparisons 
are suggested with the fossil Rhynia gwynnevaughanii, thought by someto be a highly 
organised prothallus. The book is well illustrated both with photomicrographs and well 
executed line drawings. 

A.C. JERMY 



FERNGAZ. 13(2) 1986 127 



BRITISH PTERIDOPHYTE RECORDS 

Compiled by A.J. Worland 

Since the appearance of the 'Atlas of Ferns', an annual list of additions and corrections 
has been compiled from information supplied by the Biological Records Centre, BSBI 
vice-county recorders and members of the BPS and BSBI. Lists have been published in 
the BPS 'Bulletin' up to and including 1982 and subsequently annually in the Fern 
Gazette. 

The records are presented thus: 100km square (letters are used for Irish grid 
squares to avoid confusion)/10km square followed by the recorder's name. 
Nomenclature follows the Atlas. The following additions have been received up to the 
end of March 1986. 

POST 1 980 

2.1 Lycopodiella inundata E Cumberland, R.W.M. Corner 

5.2 Selaginella kraussiana 36/67 E.P. Beattie 

7.1 Equisetum hyemale 37/41 N.F. Stewart 

7.1x2 E. x trachyodon 35/37 G. Halliday, 35/78 G.A. Swan 

7.2 E. variegatum H 12/37 F. Bonham 

7.3 E- fluviatile 53/04 N.J. Hards 

8.1 Botrychium lunaria 25/67 V. McClive 

14.1 Hymenophyllum tunbrigense 26/25 B. Simpson 

16.1 Polypodium vulgare 52/94 E.M. Hyde, 62/13 E.M. Hyde 

16.2 P. interjectum 53/05 N.J. Hards 

16.2x1 P. x mantoniae 52/94, 62/03, 62/13 E.M. Hyde 

17.1 Pteridium aquilinum 53/24 N.J. Hards 

18.1 Thelypteris thelypteroides 22/93 R.G. Woods, H 12/27 F. Bonham 

20.1 Oreopteris limbosperma 34/86 N.J. Hards, 51/04 M.D. Reed 

21.1 Asplenium scolopendrium 53/04 N.J. Hards 

21.2 A. adiantum-nigrum 43/97, 53/04, 53/14 N.J. Hards 
21.6 A. marinum 36/68 M. McAffer 

21.7a A. trichomanes subsp. trichomanes 22/72 R.N. Stringer & I.K. Morgan 

21.1 1 A. ceterach 26/40 B. Simpson 

22.1 Athyrium filix-femina 48/02 A.O. Chater, 53/34 N.J. Hards 
26.2x3 Poiystichum x bicknellii 18/52 M. Barron, 22/40 I.K. Morgan 

26.3 P. setiferum 43/97, 53/34 N.J. Hards 

27.2 Dryopteris filix-mas 53/24 N.J. Hards 

27.3 D. affinis subsp. affinis 22/40 I.K. Morgan 
27.3 D. affinis subsp. borreri 22/40 I.K. Morgan 
27.3 D. affinis subsp. stillupensis 22/72 I.K. Morgan 

27.5 D. aemula 41/22 F. Rose & R.J. Hornby 

27.6 D. villarii subsp. submontana 35/90 R.G. Jefferson 
27.8 D. carthusiana 34/85 N.J. Hards 

27.9x8 D. deweveri 22/63 I.K. Morgan, 34/85 N.J. Hards 

30.1 Azolla filiculoides 33/10 C.J. Harris, 33/20 M. Wainwright & E.D. Pugh, 

44/63 E. Chicken 



12 8 FERN GAZETTE: VOLUME 13 PART 2 (1986) 



THE FERN GAZETTE 

Original papers, articles or notes of any length on any aspect of pteridology will be 
considered for publication. 

Contributions should be sent to: 

Dr. M. Gibby, British Museum (Natural History), Cromwell Road, London SW7 5BD. 

The last date for receiving notes and articles to makethe following summer number is: 

31st December each year 

Authors should follow the general style of this number. Close adherence to the 
following notes will help to speed publication. 

NOTES FOR CONTRIBUTORS 

Manuscripts: Copy should be in English and submitted in double-spaced type with adequate 
margins, on one side of the paper only. 

Abstract. All papers, other than short notes, should include a short abstract, to beset at the head 
of the main text, indicating the scope of the topic and the main conclusions. 

Headings and sub-headings: These should follow the style of this number. (Primary sub- 
headings are centred capitals. Secondary sub-headings U and L case side roman. Tertiary sub- 
headings, if necessary are U and L case italic, ranged to the left). Numbering of sub-heading 
should be avoided. 

Latin names: Quote the authority at (usually) the first mention only, in the main text but, unless 
unavoidable for clarity not inthetitle. All latin names should be underlined throughout thetyped 
copy. 

Illustrations: Any number and combination of line and half-tone illustrations (original drawings or 
diagrams in ink, or photographs which must be black and white, and of good technical quality) 
can be included with a manuscript where these help to augment or amplify the text. Photographs 
should be of the required magnification or larger and need not be made up to full page plates. Each 
drawing or photograph should be marked on the back with details of author and figure number, 
and the top edge clearly marked "top". Illustrations will not be returned to the author unless 
specifically requested. 

Figure numbering: Grouped illustrations should follow the numbering system, fig 1a, fig 1b, fig 
2a, fig 3, etc. Figure numbers should be applied to illustrations in pencil only or on a transparent 
overlay. Final lettering will be added by the editor in a style and size consistent with the journal. 
Figure captions: Type on a separate sheet from the manuscript — include any necessary details of 
magnification as submitted (the editor will apply any correction or reduction). 

Reference lists: Please follow closely the style of this number to speed publication. Lists in 
other styles may have to be returned to authors for re-typing. 

Reprints: Twenty-five reprints are supplied free of charge to authors, who may order in advance 
further reprints which will be supplied at cost (plus postage) if requested at time of returning the 
first proofs. 



BOOKS FOR REVIEW 

Books for review in the Fern Gazette shou Id be sent to A.C. Jer my, Botany Department, 
British Museum (Natural History), Cromwell Road, London, SW7 5BD; for review in the 
Pterido/ogist books should be sent to M.H. Rickard, The Old Rectory, Leinthall Starkes, 
Ludlow, Shropshire SY8 2HP. 



BRITISH FERNS AND THEIR CULTIVARS 

A very comprehensive collection is stocked by 

REGINALD KAYE LTD 

SILVERDALE, LANCASHIRE 
CA TALOGUE ON REQUEST 



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and Plants for the cool greenhouse 

Catalogue on request 



MRS J K MARSTON 

Specialist Fern Grower 

A wide range of hardy and greenhouse ferns, especially Adiantums 

Culag, Green Lane, Nafferton, Nr. Driffield, East Yorkshire 

Send 45p for catalogue 



JACKAMOOR'S HARDY PLANT FARM 
Theobalds Park Road, Crews Hill, Enfield, Middlesex, EN2 9BG 

Tel: 01-363 4278 

Hardy ferns, unusual hardy herbaceous plants, shade and moisture loving perennials 
Send two ten-pence stamps for list 



JOIN THE EXPERTS 
Los Angeles International Fern Society 

28 page colour, illustrated journal plus fern lesson monthly, 

international spore store, educational materials and books. 

$10 domestic, $12 foreign annual dues. 

Send to 

LAIFS, 14895 Gardenhill Dr., LaMirada, CA 90638, USA 



THE NIPPON FERNIST CLUB 

There exists in Japan this large and active Society devoted 

to the study of ferns which we/comes contact with foreign pteridologists 

both amateur and professional. 

For further information write to: 

Dept. of Forest Botany, 

Faculty of Agriculture, Tokyo University, Hongo, 

Bunkyo-ku, Tokyo, Japan 113 



The British Pteridological Society 
THE FERN GAZETTE 

VOLUME 13 PART 2 1986 



CONTENTS 



Page 



MAIN ARTICLES 

The modular growth of Lycopodium annotinum 

- Terry V. Callaghan, Brita Svensson & Alistair Headley 65 

The ecology of pteridophytes in Tasmanian cool temperate rainforest 

- S.J. Jar man. G. Kantvilas & M.J. Brown 11 

Tmesipteris in Vanuatu (New Hebrides) 

- A.F. Braithwaite 87 
Effects of salinity on gametophyte growth of A crost /chum aureum 

and/4, danaeifolium 

- Robert M. Lloyd & Donald P. Buckley 97 

The ecology of pteridophytes in the Mwanihana Forest Reserve, Tanzania 

- J.C. Lovett & D.W. Thomas 103 
George Gardner's pteridophyte herbarium and lectotypes of Gardner's 

new fern species 

- B.S. Parris 109 

A new Dryopteris hybrid from Spain 

- Christopher R. Fraser-Jenkins & Mary Gibby 1 1 3 

Subgeneric names in Selaginella 

- A.C. Jermy 117 

SHORT NOTES 

The occurrence of Schizaea dichotoma in Tanzania 

- J.C. Lovett 119 
A chromosome count for Anogramma leptophylla in Madeira 

- Mary Gibby 1 20 

Cystopteris in the Cape Verde and Canary Islands 

- Wolfram Lobin 121 

Cystopteris viridu/a in mainland Africa 

- C.R. Fraser Jenkins 121 

First record of Equisetum x dycei in continental Europe 

- H. W. Bennert & U. Peters 1 22 
An anomaly of Botrychium matricariifo/ium 

- Harald Schumann 1 23 

Corrections to Index Filicum 

- B.S. Parris 123 

Obituary: Professor E.A.C.L.E. Schelpe 

- A. V. Hall 1 24 

REVIEWS 1 08, 1 1 2, 1 1 6, 1 25, 1 26 

BRITISH PTERIDOPHYTE RECORDS 127 

(THE FERN GAZETTE Volume 13 Part 1 was published on 29th August 1985) 

Published by THE BRITISH PTERIDOLOGICAL SOCIETY, c/o Department of 
Botany, British Museum (Natural History), London SW7 5BD 



ISSN 0308—0838 

Metloc Printers Ltd.. Old Station Road, Loughton. Essex 



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GAZETTE 



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Edited by 

J.A£rabbe,M.6ibby 
& BSParris 



3R1TISH 

PTER1D0L0G1CAL 
SOCIETY 



Volume 13 Part 3 



J 



1987 



THE BRITISH PTERIDOLOGICAL SOCIETY 
Officers and Committee for 1987 

President: G. Tonge 
President Emeritus: J.W. Dyce 

Vice-Presidents: Dr R.E. Holttum, F. Jackson, A.C. Jermy, 

R. Kaye, Prof. I. Manton 

Honorary General Secretary and A.R. Busby, 'Croziers', 

Editor of the Bulletin: 16 Kirby Corner Road, Canley, Coventry, CV4 8GD 

(Tel: Coventry 715690) 

Assistant Secretary (Membership): A.M. Paul, 

Department of Botany, British Museum (Natural History), 
Cromwell Road, London, SW7 5BD 

Treasurer: Dr B.A. Thomas, Botany Department, National Museum of Wales, 

Cathays Park, Cardiff, CF1 3NP 
assisted by Dr N.J. Hards 

Meetings Secretary: Vacancy, enquiries to A.R. Busby 

Editors of the Fern Gazette: J.A. Crabbe, Dr M. Gibby, Dr B.S. Parris 

Material for publication should be sent to Dr B.S. Parris, 
Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AE 

Editor of the Pteridologist: M.H. Rfckard, The Old Rectory, Leinthall Starkes, 

Ludlow, Shropshire, SY8 2HP 
assisted by J.W. Dyce 

Committee: J. Bouckley, J.M. Camus, C.R. Fraser-Jenkins, Dr N. Hards, 

J. Ide, R.P.H. Lamb, A.M. Paul, A. Pigott, P. Ripley, Dr A. Willmot 

Fern Distribution Recorder: A.J. Worland, 102 Queens Close, Harston, 

Cambs., CB2 5QN 

Spore Exchange Organiser: R.F. Cartwright, 13 Perry Mill Road, Peopleton, 

Pershore, Worcestershire 

Archivist: N.A. Hall, 15 Mostyn Road, Hazel Grove, Stockport, Cheshire, SK7 5HL 

Booksales Organiser: J.W. Dyce, 46 Sedley Rise, Loughton, Essex IG10 1LT 

Trustees of Greenfield Fund: G. Tonge, A.R. Busby, Dr B.A. Thomas 

The BRITISH PTERIDOLOGICAL SOCIETY was founded in 1 891 and today continues as a focus for 
fern enthusiasts. It provides a wide range of information about ferns through the medium of its 
publications and available literature. It also organises formal talks, informal discussions, field 
meetings, garden visits, plant exchanges and a spore exchange scheme. The Society has a wide 
membership which includes gardeners, nurserymen and botanists, both amateur and professional. 
The Society's journals, the Fern Gazette, Pteridologist and Bulletin, are published annually. The Fern 
Gazette publishes matter chiefly of specialist interest on international pteridology, the Pteridologist, 
topics of more general appeal, and the Bulletin, Society business and meetings reports. 

Membership is open to all interested in ferns and fern-allies. SUBSCRIPTION RATES (due on 1st 
January each year) are Full Personal Members £ 10; Personal Members not receiving the Fern 
Gazette £7.50; Student Members £5; Subscribing Institutions £12. Family membership in any 
category is an additional £2. Applications for membership should be sent to the Assistant Secretary 
(address above) from whom further details can be obtained. (Remittances made in currencies other 
than Sterling are £1.50extra to cover bank conversion charges). Airmail postage for all journals — an 
extra £3.50, or for those not receiving the Fern Gazette £2.50. 



Back numbers of the Gazette, Pteridologist and Bulletin are available for purchase from the 
Treasurer (address above), from whom further details can be obtained. 



FERN GAZ. 13(3) 1987 



129 










This number is dedicated to 

PROFESSOR DOCTOR TADEUS REICHSTEIN, 

eminent pteridologist, 

on the occasion of his Ninetieth Birthday 

20th July 1987 



Photograph courtesy of H. & K. Rasbach 



130 FERN GAZETTE: VOLUME 13 PART 3 (1987) 



Further papers submitted to honour Professor Reichstein 
will appear in the next number of the Fern Gazette 



FERN GAZ. 13(3) 1987 131 



T. REICHSTEIN : A PERSONAL APPRAISAL 

There can be no doubt that, in Tadeus Reichstein, now entering his 90th year, we have 
among us one of the most outstanding personalities of the 20th century. I am not 
erudite enough to discuss his chemical achievement perse except to note that in 1968 
he was awarded the highest accolade in the gift of the Royal Society of London (the 
Copley Medal). Other aspects of his life and scientific interests can perhaps best be 
introduced in an anecdotal manner based on my own personal experience of how I first 
became involved with him. 

This happened early in the 1950s when he wrote to me from the Chemistry 
Department in Basel to follow up some information that I had recently published in my 
book (Problems of Cytology and Evolution in the Pteridophyta, 1 950, CUP). This book had 
been drawn to his attention by a Swiss colleague and he wanted to know how best to 
confirm the putative hybrid nature of some of his local ferns. In reply, I offered to visit 
him to look at them provided only that he could lay on any kind of a student's 
microscope. He replied by telegram saying "Splendid. Come this weekend". This 
message reached me at 12 noon on Whit Saturday and if one can effectively think back 
to that time there will be no doubt regarding the impossibility of flying to Switzerland at 
Whitsun if you don't know that you want to go before noon on Whit Saturday. I had to 
explain this also by telegram but having secured a flight as soon as possible at a later 
weekend (in term time I could not easily be absent from Leeds except at a weekend) I 
took the precaution, before setting off, of consulting our then professor of organic 
chemistry for information about my unknown correspondent. The reply I received was 
surprising, and at first alarming. I was told "Oh, Reichstein. He is a very well known 
mountain climber with many difficult traverses named after him". This rather 
frightening prospect was toned down by one additional fact, namely that he was now 
old and therefore might not want to dangle me on a rope over a precipice. Somewhat 
comforted by that I set off and was received with the most incredible hospitality and 
kindness. I had no difficulty in sorting out his problematical hybrids by showing him 
how to recognise bad spores under a microscope. I also took the opportunity of taking 
fixings of several plants that he had growing in his greenhouse from local gatherings 
and some of these, when looked at chromosomally after my return to Leeds proved to 
be new — diploids where tetraploids were to be expected or the converse. Herewith 
began an active collaboration that has lasted until the present day. 

Even at the first encounter I was so much impressed with his ready intelligence as 
well his botanical expertise that I made more enquiries as soon as I got back to Leeds, 
this time from a student in the Chemistry Department. This lad's eyes opened wide as 
soon as I mentioned the name Reichstein. I was assured that he was a tremendous 
chemist and indeed the holder of a Nobel Prize (awarded in 1 950). What a difference 
from the previous concentration on mountain climbing as a major credential, true as 
this doubtless is! 

To a professional chemist, the very simple reagents needed (acetic acid and 
absolute alcohol) to take fixations for aceto-carmine squash preparations offer no 
problems and I was soon receiving new fixations of additional Swiss specimens. At 
first I dealt with these myself but soon began to pass them onto John Lovis (a member 
of my staff) who benefitted greatly by the experience. Since Reichstein himself owned 
a powerful car and was willing to apply his alpine expertise in any part of Europe 
(France, Spain, Germany, Italy etc.) as required, knowledge of European ferns rapidly 
increased. Certain key genera, notably Aspleniu m, Polypodium, Dryopteris etc., are 
now as fully known as they are likely to become, at least with respect to this part of the 
world. For this alone, I was able, in 1974, to propose Reichstein as an honorary foreign 
member of the Linnean Society of London, when I became president (1 974-76) of that 
well known biological society. 



13 2 FERN GAZETTE: VOLUME 13 PART 3 (1987) 



Reichstein has now been able to stimulate his own local group to become experts 
both in cytology and in taxonomy thereby virtually eliminating any former dependence 
on Britain. Moreover, the Swiss group has recently turned the tables on Britain by 
giving invaluable help towards finalising a long drawn out project on the fern flora of 
Madeira begun by the Leeds group in 1949 but published only in 1986 (see Bull. Brit. 
Mus. (Nat. Hist.) Bot. 15: 123-1 61). By that time four authors had necessarily become 
involved namely I. Manton (Leeds), John Lovis (now permanently resident in New 
Zealand), G. Vida (Budapest) and Mary Gibby (London). Such a geographical spread 
among authors posed a major obstacle to communication and the text took two years to 
complete. Indeed, without Reichstein's constant help in co-ordinating nomenclature 
and literature and in detecting errors sometimes involving the typescripts of one or 
other author finality might never have been reached. 

A pioneer as effective as this in so many different fields of human activity is rare 
indeed and anyone privileged to have known him personally can scarcely fail to be both 
proud and grateful for the experience. 

Irene Manton 



FERN GAZ. 13(3) 1987 133 



ASPLENIUM X REICHSTEINII (= ASPLENIUM FONTANUM X 

A. MAJOR/CUM; ASPLENIACEAE: PTERIDOPHYTA), 

A NEW ENDEMIC FERN HYBRID FROM MALLORCA, 

BALEARIC ISLANDS 



H. WILFRIED BENNERT 

Spezielle Botanik, Fakultat fur Biologie, Ruhr-Universitat Bochum, 

Universitatsstrasse 150, D-4630 Bochum 1, Federal Republic of Germany 

HELGA RASBACH and KURT RASBACH 
Datscherstrasse 23, D-7804 Glottertal, Federal Republic of Germany 

ABSTRACT 

An Asplenium hybrid that was found near the town of Soller, Mallorca, is 
described and the name Asplenium x reichsteinii Bennert & Rasbach is proposed 
for it. It is a triploid plant exhibiting 36 bivalentsand36 univalents at meiosis. These 
cytological results as well as its morphology strongly suggest that it originated from 
a cross between Asplenium fontanum and/4, majoricum. Further support in favour 
of this interpretation was obtained by comparing it with hybrid plants of this 
combination artificially produced by Sleep(1 967); an almost complete agreement in 
morphology and an identical cytological pattern were both observed. The 
occurrence of Asplenium fontanum on Mallorca and earlier reports in literature of a 
single plant obviously also representing Asplenium x reichsteinii are discussed. 

INTRODUCTION 
Asplenium majoricum Litard. is an endemic fern of the Island of Mallorca where it 
occurs at only a few scattered localities, most of them in the close vicinity of Soller 
(Jaquotot & Orell 1968). It is an allotetraploid species (Jermy & Lovis 1964) having 
originated from a cross between diploid Asplenium petrarchae subsp. biva/ens (D.E. 
Meyer) Lovis & Reichstein and another diploid, Asplenium fontanum (L.) Bernh., by 
chromosome doubling (Sleep 1967, 1983; Lovis & Reichstein 1969; Lovis etal. 1 969). 
Lovis (1977; p. 326) succeeded in resynthes\z'mg Asplenium majoricum from its diploid 
parents. 

Obviously Asplenium major/cum hybridizes with other species of Aspleniaceae 
rather easily. Three described and named hybrids involving Asplenium majoricum are 
presently known, all of them found near Soller: Asplenium majoricum x A. 
trichomanes subsp. quadrivalens D.E. Meyer (= Asplenium x ore/Hi Lovis & Reichstein, 
Lovis & Reichstein 1 969), Asplenium majoricum xA. petrarchae (Guerin) DC. subsp. 
petrarchae {- Asplenium x sollerense, Lovis, Sleep & Reichstein, Lovis etal. 1969), and 
finally the intergeneric hybrid Asplenium majoricum x Ceterach officinarum DC (= x 
Asp/enoceterach barrancense Bennert & D.E. Meyer, Bennert & Meyer 1972). 

During an excursion to the olive groves above Biniaraix near Soller, Mallorca, in 
April 1 986 two plants were found that were believed to represent the cross between 
Asplenium fontanum and A. majoricum. The area was revisited in October 1 986 and 
another four hybrid plants were discovered. Results of further studies, especially the 
cytological examinations, confirmed the assumed origin of this hybrid. It is described 
here in detail and named Asplenium x reichsteinii. 

MATERIALS AND METHODS 
The two plants of Asplenium x reichsteinii found in April 1986 were collected and 
taken into cultivation in the greenhouse of Prof. Reichstein at Basel and at Bochum for 
further studies. In October 1986 another plant (out of a group of three plants growing 
closely together) was taken into cultivation at Basel. Of the fourth plant found in 



134 



FERN GAZETTE: VOLUME 13 PART 3 (1987) 



October 1 986 fixations were made in the field. The meiosis was investigated applying 
the classical acetocarmine method (Manton 1950). For the final analysis and the 
photographs of the spore mother cells a magnification of 1,000 x and a phase 
microscope were used. In the case of the type plant 18 cells inthestageofdiakinesis/ 
metaphase 1 were counted. 

DIAGNOSIS 
Asplenium x reichsteinii Bennert et Rasbach hybr. nov. (= Asplenium fontanum (L.) 
Bernh. x Asplenium major/cum Litard.). 

Planta hybrida, media inter parentes, sed Asplenio fontano aliquid similior. Ab hoc 
distinguitur: pinnae magis compactae, minus profunde incisae. Sporae abortivae. Planta 
triploidea, meiosi chromosomatibus bivalentibus 36, univalentibus item 36. 

Holotype: Asplenium x reichsteinii Bennert et Rasbach. On a limestone wall above 
Biniaraix near Soller, Mallorca, Balearic Islands, Spain; c. 180m alt..: with Asplenium 
majoricum, A. petrarchae, A. trichomanes, and Ceterach officinarum growing in the near 
vicinity: leg.: H.W. Bennert et U. Peters, 1 7 April 1 986; WB SP43/86; plant later cultivated 
in Basel under reference number TR-6477: whole plant pressed on 7 October 1986; 
holotype: B (see Fig. 1 ). 

Dedicated to Prof. Dr. T. Reichstein, Basel, who has added very much to the knowledge of 
the ferns, especially of the genus Asplenium. 




FIGURE 1 .Asplenium x reichsteinii Bennert et Rasbach (= Asplenium fontanum xA majoricum); 
silhouette of the type plant (kept in B). 



ASPLENIUM X REICHSTEINII 



135 



r5 




cm 






FIGURE 2. Asplenium x reichsteinii, its parents and the artificially produced hybrid A splenium 
major/cum xA. fontanum; silhouettes of fronds; a: Asplenium fontanum; Gorge duVerdon, Dept. 
Var, France; WB 52/72, 3.4.1972; b,c: Asplenium x reichsteinii; Biniaraix near Soller, Mallorca, 
Spain; Ras-535, 13.10.1986; d: Asplenium majoricum; Puig d'en Barrera near Soller, Mallorca, 
Spain; WB 167/71, 10.4.1971; e, f: Asplenium x reichsteinii; Biniaraix near Soller, Mallorca, 
Spain; WB 46/86, (= SP 24/86) 1 7.4 1 986; g, h: Synthesized hybrid between Asplenium majoricum 
o. (Soller, Mallorca x A. fontanum cf (Roche, Switzerland; AS 266 (i), 30.5.1962. The arrows 
indicate the position where the colour of the rachis or of the stipe changes from brown below to 
green above. As this position is different on both surfaces solid arrows were used as markets for 
the upper surface and broken arrows for the lower surface. 



!36 FERN GAZETTE: VOLUME 13 PART 3 (1987) 



Paratypes: A second plant of Asplenium x reichsteinii was discovered on the same day 
(17.4.1986; leg.: H.W. Bennert et U. Peters) somewhat above (c. 200m alt.) the locality of 
the holotype. This plant is in cultivation in Bochum (as SP 24/86); fronds of it will be 
deposited in the following herbaria: BM, G, K, MA and Z. A third plant collected on 13 
October, 1 986 (leg.: H.W. Bennert, H. Rasbach et K. Rasbach) was divided into two parts 
both being cultivated in Basel (as TR-6540 a & b). Of a fourth plant (that remained in the 
field) fixations were made and fronds were collected on 13 October, 1986; these are kept in 
the private herbaria of H. and K. Rasbach (Ras-535) and H.W. Bennert (WB 71 /86). The 
cytological examinations that were made of all paratype plants gave the same results as in 
the case of the holotype. 

Hybrid plant with its gross morphology being intermediate between both parents, 
however rather similar to Asplenium fontanum from which it can be distinguished by the 
following characters. Pinnae more compact and less deeply cut: pinna segments less 
sharply toothed. Stipe and lower part of the rachis (especially on the abaxial surface) dark 
brown coloured. Scales of rhizome in their colour intermediate between the parent species 
(in Asplenium fontanum light brown, in Asplenium majoricum deep brown). Fronds (of the 
wild growing plants) up to 12cm long and 1.5cm broad. Spores abortive (Fig. 3). Plant 
triploid with 36 bivalents and 36 univalents at meiosis. 





TV 

[mm 



FIGURE 3. Ripe sporangia of Asplenium x reichsteinii containing aborted spores. 

CYTOLOGY AND ORIGIN OF ASPLENIUM X REICHSTEINII 
The morphology of Asplenium x reichsteinii clearly suggests a relationship to 
Asplenium fontanum andAsp/enium majoricum (see Fig. 1 and Fig. 2). Further support 
for this interpretation comes from the cytological results. As Asplenium x reichsteinii 
is triploid it must have originated from a cross between a tetraploid and a diploid 
species. The pairing behaviour of chromosomes showing 36 bivalents and the same 
number of univalents at meiosis (Fig. 4) can be interpreted in two different ways. One 
possibility is that an autotetraploid (AAAA) and an unrelated diploid species(BB) were 
involved with the bivalents originating from the autotetraploid species exclusively 
(AAB). The same pairing behaviour may result, however, if an allotetraploid species 
(AABB) and a diploid species containing a related genome (AA or BB) hybridize; in this 
case each species contributes one set of chromosomes to the bivalents observed at 
meiosis (AAB or ABB). 

Considering the first possibility Asplenium trichomanes, A. petrarchae and A 
ruta-muraria are the autotetraploid species, Asplenium fontanum (see following 
chapter) and A. onopteris L. the diploid ones that occur in the vicinity of Soller thus 
representing possible candidates involved in hybridization. None of the hybrid 



ASPLENIUM X REICHSTEINII 



137 



.. •> • 






< 



»> 









^ 



10yur 






^ 



cO 



<& 



£ 

£?■• 



4**- 

r S8," 

<? 



„ ,, '-** 






^ 

U 



etf\ 



lOfir 



FIGURE 4. Cytology oiAsplenium x reichsteinii (type plant WB SP 43/86); a: Photograph of spore 
mother cell during meiosis (diakenesis) showing 36 bivalents and 36 univalents; b: Explanatory 
diagram; bivalents black, univalents in outline; preparation by H.R. 



13 8 FERN GAZETTE: VOLUME 13 PART 3 (1987) 



combinations between these species would, however, result in a plant exhibiting such 
an Asplenium fontanum-\\ke morphology as Asplenium x reichsteinii does. Even 
hybrids involving Asplenium fontanum should show closer morphological 
relationships to the tetraploid parent as the latter would contribute two sets of 
chromosomes which would dominate over the single set coming from A. fontanum. 

When checking the second possibility of explaining the cytological results only 
one allotetraploid species has to be considered, namely Asplenium majoricum. 
Keeping in mind that it contains two genomes of Asplenium fontanum and A. 
petrarchae (FoFoPePe, see introduction) only a cross involving A splenium fontanum 
(FoFo) or diploid A. petrarchae (PePe) would show the pairing behaviour observed. 
Apart from the fact that diploid Asplenium petrarchae is to date not known to occur on 
the Balearic Islands the hybrid involving this species (FoPePe) should be close to A 
petrarchae in its morphology. The only hybrid combination to remain is that between 
Asplenium majoricum and A. fontanum. This hybrid must be triploid and have the 
genome formula FoFoPe with one genome Fo being derived directly from the diploid 
parent Asp/en ium fontanum and the other one being contributed by A majoricum. 
This indeed explains both the morphological features (two genomes Fo dominate over 
one Pe) as well as the cytology (the bivalents being formed by the two genomes Fo) of 
Asplenium x reichsteinii. 

Though this interpretation of the origin of Asplenium x reichsteinii hardly can be 
doubted it is, for reasons that will become obvious in the following chapter, of 
considerable value to obtain further support from independent investigations. In order 
to elucidate the origin and interrelationships of Asplenium majoricum. Sleep (1967) 
produced hybrids in various combinations. Within her hybridization programme she 
successfully synthesized the hybrid between Asplenium fontanum and A majoricum. 
For this plant she reports exactly the same pairing behaviour of chromosomes as 
observed in Asplenium x reichsteinii. Furthermore, the morphology of this artificially 
produced hybrid is almost identical with that of Asplenium x reichsteinii from Mallorca 
(Fig. 2). The only obvious difference is that in the extension of the brown colour of the 
rachis (which especially in hybrids may vary to some extent). The two fronds of the 
synthesized hybrid that are shown in Fig. 2, g and h, will be deposited in B together 
with the holotype. 

ON THE OCCURRENCE OF ASPLENIUM FONTANUM ON MALLORCA 
Asplenium majoricum as one parent species of Asplenium x reichsteinii is rather 
frequent in the area where the hybrid plants were found. To our surprise not even a 
single plant of Asplenium fontanum could be discovered although several days were 
spent investigating the fern flora around Soller with special attention given to the 
occurrence of this species. In modern literature Asplenium fontanum is consistently 
listed as a rare but undoubted member of the Majorcan flora (Jalas & Suominen 1 972, 
Bonafe 1977, Duvigneaud 1979, Pichi Sermolli 1979, Greuter et al. 1981, Bolos & 
Vigo 1 984, Salvo et al. 1 984, Smythies 1 984, Castroviejo et al. 1 986). Nevertheless, 
any authentic and reliable recent report that would confirm the present occurrence of 
Asplenium fontanum on Mallorca seems to be lacking. Judging from the information 
given by Bonafe (1 977) all corresponding original reports date back to the time before 
1920. The oldest source that mentions Asple n ium fontanum from Mallorca seems to 
be the catalogue of vascular plants of the Balearic Islands by Mares & Vigineix(1 880). 
At that time Asplenium majoricum had not yet been described as a separate species; 
this was done by Litardiere in 1911. Therefore it seems doubtful whether really 
Asplenium fontanum and not/4, majoricum was meant. In his chapter onAsp/enium 
fontanum Bonafe (1977) also mentions the existence of herbarium specimens and 



ASPLENIUM X REICHSTEINII 



139 



presents a photograph (p. 34) showing a complete dried plant being kept in the "Herbari 
Bianor de Palma". This voucher could indeed belong XoAsplenium fontanum. Bonafe 
(1 977) further reports that specimens obviously belonging to the same collection are 
kept in the herbarium of the Botanical Institute at Barcelona. The sheet with these 
specimens (no. 72095) was borrowed from Barcelona and thoroughly examined. On 
the basis of morphology (shape of pinnae, green colour of stipe and rachis) and the well 
developed (not aborted) spores with an exospore length of (29-)32-35(-41 )um it is 
obvious that these plants indeed represent true Asplenium fontanum (Fig. 5). The 




FIGURE 5. Herbarium specimen of Asplenium fontanum from Soller, Mallorca, kept in the 
herbarium of the Botanical Institute at Barcelona (no. 72095). Leg.: F.re Bianor-Marie; 
15.11.1917. Silhouette of whole plant. 



inscription on the label is as follows: "Plantes des Baleares Asplenium Halleri, R.Br. 
Murs des olivaies. Soller, Est. 1917, 15 Novembre. F.re Bianor-Marie". This find was 
obviously the basis for the report of Asplenium Halleri (a synonym for Asplenium 
fontanum) in the plant list of Bianor (1917). Although he does not mention/lsp/e/?/t//77 
majoricum (which by that time had already been described as a separate species) 
Bianor's statement must be accepted as correct. More recently collected specimens in 
the private herbarium of J. Orell, Palma de Mallorca, with a morphology close to 
Asplenium fontanum were checked and found to belong XoAsplenium x reichsteinii 
(see following chapter). 

If Asplenium fontanum still exists on- Mallorca it must be a very rare species 
possibly restricted to one or very few localities, with only a small number of individuals. 
We are inclined to believe that it would not primarily grow on the walls of the olive 
terraces as Asplenium majoricum and/4, petrarchae do but prefer more favourable 
microhabitats on limestone cliffs. These should be shady and moist to ensure a 
relatively good water supply especially during summertime when long lasting rainless 
periods appear which are typical of the Mediterranean climate. Such habitats are more 
likely to occur at higher elevations in the mountains where as a consequence of 
frequent cloud formation air humidity tends to be high and water may condensate thus 
improving plant water relations. Asplenium fontanum is certainly less drought 



140 FERN GAZETTE: VOLUME 13 PART 3 (1987) 



resistant than Asplenium petrarchae and A. majoricum or other members of this 
genus. 

Although we could not establish the present occurrence oi Asplenium fontanum 
on Mallorca there is good reason to believe that it was, and is, still there. One argument 
is that Asplenium fontanum is one parent species of A. majoricum and it is most 
probable that the latter originated on Mallorca where it is endemic. The second point is 
the rather frequent occurrence oi Asplenium x reichsteinii (see following chapter). The 
normal situation for fern hybrids to arise is that both parent species are present often 
growing closely together. The tendency of Asplenium x reichsteinii to occur at 
somewhat higher elevations (around 200m alt., highest locality at 235m alt.) and its 
preference for natural limestone cliffs (only 2 out of 6 plants grew on walls), always at 
the base of steep and high projecting rock faces, are remarkable and may be indicators 
for the habitats where Asplenium fontanum can be expected on Mallorca. 

EARLIER REPORTS ON HYBRIDS BETWEEN ASPLENIUM FONTANUM AND A. MAJORICUM 
The oldest published report on hybrids between Asplenium fontanum and A, 
majoricum seems to be that by Sleep (1 967). As already mentioned she produced this 
cross artificially within a hybridization programme to study the origin of Asplenium 
majoricum. Under the applied experimental conditions it formed rather easily (6.4% 
success). 

Besides these artificially raised hybrids one wild plant that was found near Soller by 
Schulze, Orell and Bonafe in 1964 (det. 24. III. 1964) obviously represents the cross 
between Asplenium fontanum and A. majoricum. This plant is referred to in several 
publications (Jaquotot & Orell 1968, Lovis & Reichstein 1969, Bonafe 1977, 
Reichstein 1 981 , Castroviejo et al. 1 986) but without having received a valid name. 
Reichstein who checked pressed fronds of it (kept in the private herbarium of J. Orell, 
Palma de Mallorca) did not come to a final conclusion in his earlier paper (Lovis & 
Reichstein 1969) where he considers it asAsp/enium fontanum or the hybrid between 
A. fontanum and A. majoricum. Later (Reichstein 1 981 ), however, he stated that the 
assignment of Jaquotot & Orell (1968) (Asplenium fontanum xA. majoricum) is most 
probably correct. The fronds in the herbarium of Orell were checked again in 1 986 by 
the present authors and only sporangia with abortive spores were found. It is therefore 
obvious that this plant indeed represents Asplenium x reichsteinii. Very recently 
Antoni Rebassa, a biology student living in Soller, sent 2 fronds of another plant of 
Asplenium x reichsteinii collected in October 1986. He describes the locality where he 
found the hybrid as follows: "Son Vencis", Ses Tres Creus, Soller. This is probably the 
same area where Schulze, Orell and Bonafe found their plant in 1964. 

This means that until now at least 8 different individuals of Asplenium x 
reichsteinii have been found in nature. As the hybrid resemb\esAsp/enium majoricum 
it may easily be overlooked unless it is examined closely. Therefore more hybrid plants 
are to be expected and may be discovered in the neighbourhood of Soller. 

ACKNOWLEDGEMENTS 
We thank Dr A. Sleep, Leeds, for providing us with two very valuable fronds of the 
artificially produced hybrid between Asplenium fontanum and A majoricum, Mr J. 
Orell, Palma de Mallorca, for letting us examine his specimen of Asplenium x 
reichsteinii, Prof. DrT. Reichstein, Basel, for careful cultivation of the type plant and Dr 
J. Montserrat, Barcelona, for sending us the voucher of Asplenium fontanum from 
Mallorca on loan. Furthermore we gratefully acknowledge the help of Prof. Dr H. 
Haeupler, Bochum, who not only gave some information on floristic literature of the 
Balearic Islands but also enabled one of us (W.B.) to spend two days in the Soller area 
during a students' excursion to Mallorca in April 1 986. Our thanks are extended to Dr 



ASPLENIUM X REICHSTEINII 141 



B.S. Parris, Kew, Prof. Dr K.U. Kramer and Dr J.J. Schneller, both Zurich, for reviewing 
the manuscript and correcting our English; Prof. Dr K.U. Kramer also improved the 
Latin diagnosis. We are also grateful to Mrs I. Kiinzel, Bochum, who assisted us with 
the photographic work. 

REFERENCES 

BENNERT, W. & MEYER, D.E. 1972. Der GaXXungsbastardx Asplenoceterach barrancense hybr. 

nov. (Asplenium majoricum Lit. xCeterach officinarum Lam. et DC). Willdenowia 6: 461 - 

470. 
BIANOR, EC. 1917. Plantes de Mallorca. But//. Inst. Catal. Hist. Nat. XVII, 133-150. 
BOLOS, 0. de & VIGO, J. 1 984. Flora dels Paisos Catalans. Volum I (Introduccio. Licopodiacies - 

Capparacies). Barcelona. 
BONAFE, F. 1977. Flora de Mallorca. Volum I. Palma de Mallorca. 
CASTROVIEJO, S., LAINZ, M., LOPEZ GONZALES, G., MONTSERRAT, P., MUNOZ GARMENDIA, 

F., PAIVA, J. & VILLAR, L. (Eds.) 1 986. Flora iberica. Plantas vascu/ares de la Peninsula 

Iberica e Is/as Ba/eares. Vol. I Lycopodiaceae-Papaveraceae. Madrid. 
DUVIGNEUAD, J. 1979. Catalogue provisoire de la f/ore des Ba/eares. Soc. pour I'echange des 

plantes vase, de I'Europe occ. et du bassin medit. 7 /, suppl. Liege. 
GREUTER, W., BURDET, H.M. & LONG, G. (Eds.) 1981 .Med-Checklist. I. Pteridophyta. Geneve & 

Berlin. 
JALAS, J. & SUOMINEN, J. (Eds.) 1 972. Atlas Florae Europaeae. Vol. 1 Pteridophyta (Psilotaceae 

to Azollaceae). Helsinki. 
JAQUOTOT, C. & ORELL, J. 1968. Asplenium majoricum R. Litardiere; suarea de expansion en la 

sierra norte de Mallorca. Collectanea Bot. 7: 559-571. 
JERMY, A.C. & LOVIS, J.D. 1 964. Asplenium majoricum Litardiere. Brit. Fern Gaz. 9: 163-167. 
LITARDIERE, M.R. de 1911. Contribution a I'etude de la flore pteridologique de la peninsule 

iberique. Bull. Geogr. Bot. 21: 12-30. 
LOVIS, J.D. 1977. Evolutionary patterns and processes in ferns. Adv. Bot. Res. 4: 229-415. 
LOVIS, J.D. & REICHSTEIN, T. 1969. Der Fambastard Asplenium x ore/lii hybr. nov.= Asplenium 

majoricum Litard. x A. trichomanes L. subsp. quadrivalens D.E. Meyer und die 

Abstammung von A. majoricum. Ber. Schweiz. Bot. Ges. 79: 335-345. 
LOVIS, J.D., SLEEP, A. & REICHSTEIN, T. 1969. Der Fambastard Asplenium x sollerense hybr. 

nov. = Asplenium majoricum Litard. xA petrarchae (Guerin) DC. subsp. petrarchae. Ber. 

Schweiz. Bot. Ges. 79: 369-376. 
M ANTON, I. 1 950. Problems of cytology and evolution in the Pteridophyta. Cambridge. 
MARES, P. & VIGINEIX, G. 1 880. Catalogue raisonne des plantes vasculaires des lies Ba/eares. 

Paris. 
PICHI SERMOLLI, R.E.G. 1 979. A survey of the pteridological flora of the Mediterranean Region. 

Webbia34: 175-242. 
REICHSTEIN, T. 1981. Hybrids in European Asp/eniaceae (Pteridophyta). Bot. Helv. 91: 89-139. 
SALVO, A.E., CABEZUDO, B. & ESPANA, L. 1 984. Atlas de la pteridoflora iberica y balear./lcfa 

Bot. Malacitana 9: 105-128. 
SLEEP, A. 1 967. A contribution to the cytotaxonomy of Asplenium majoricum. Brit. Fern Gaz. 9: 

321-329. 
SLEEP, A. 1983. On the genus Asplenium in the Iberian peninsula. Acta Bot. Malacitana 8: 1 1 - 

46. 
SMYTHIES, B.E. 1984. Flora of Spain and the Balearic Islands. Checklist of Vascular Plants. I. 

Pteridophyta, Gymnospermae, Acanthaceae-Crassulaceae. Englera 3 (1): 1 -212. 



14 2 FERN GAZETTE: VOLUME 13 PART 3 (1987) 



REVIEW 

INDEX OF THELYPTERIDACEAE by J.W. Grimes and B.S. Parr is. 50 pp. 154x 
243mm. Royal Botanic Gardens, Kew. ISBN 0947643 03 6. 1986. Price £6.30 
(over the counter) or £7.25 find. p. & p) from Dr B.S. Parr is at R.B.G., Kew, 
Richmond, Surrey, TW3 3AE. 

This slender book consists of an easily referable index, in the form of three columns 
across the page. The first column is an alphabetical arrangement of all the basionyms 
of species which are now included in the Thelypteridaceae. Column two gives the 
genus, author and date in which that epithet was originally described: column three 
gives the present (or proposed) disposition, with author anddate when published, or in 
the case of synonyms, the accepted species name, which may then be looked up in 
column one. For full bibliographical references one must refer to Index Filicum; all 
modern taxonomic treatments, up to the end of 1985, have been covered. 

Thelypteridaceae is a large and confusing group and such an index cannot befinal. 
We are promised alterations and additions in the future in the form of Supplements. As 
pointed out in the Introduction, opinions will differ and there are doubtless some 
mistakes or omissions. I am sure they will be few. This little book, well bound in a soft, 
but durable, plastic-impregnated cover, is a sensible and very useful spin-off of 
herbarium curation. 

A.C. JERMY 



FERN GAZ. 13(3) 1987 



143 



NOTES ABOUT ASPLENIUM I. ASPLENIUM QUEZELII 

A PSEUDO-ENDEMIC SPECIES IDENTICAL WITH 

A. DAGHESTANICUM (ASPLENIACEAE : PTERIDOPHYTA) 



R.L.L. VIANE 

Leerstoel voor Morfologie, Systematiek en Ecologie der Planten, 

State University of Gent, K.L. Ledeganckstr. 35, B-9000 Gent, Belgium 

ABSTRACT 

A comparative morphological study has shown that A. quezelii and A. 
daghestanicum are identical. A daghestanicum is a relic (Caucasian) element of the 
Tibesti flora, with close relatives in the Himalayas and China. 

With great pleasure I dedicate this article to Prof. T. Reichstein. We have been working 
together to get a better understanding of various "difficult" Asplenia. Reichstein's 
interest in this genus is illustrated by the 27 articles he wrote about it in the last 25 
years. The discovery of some new species in the Himalayas and China, similar to 
Asplenium daghestanicum Christ and/4. <7tyeze//7Tardieu-Blot, renewed our interest in 
this group. Several new species await description. This paper only deals with the 
identity of A. daghestanicum and A. quezelii. 

INTRODUCTION 
H. Christ described A. daghestanicum in 1906 from a single collection (5 plants) made 
by Alexcenko and Woronow (holotype : P !, isotype : BR !). The ferns were collected in 
1902 near Kurag in Daghestan (NW Caucasus). Christ suggested some distant 
relationship with A. fontanum Bernh., but/4, daghestanicum became almost forgotten 
until Reichstein et al. (1 973) compared it with A. creticum Lovis, Reichst. & Zaffran. At 
Reichstein's request Askerov collected the species again, in the same area, in 1 982. 
Spores of this collection were used to raise progeny (TR -6062) for cytological studies 
(Askerov et al. in prep.). Until now, A daghestanicum was considered to be an endemic 
of Daghestan. 

In 1956 P. Quezel collected the single plant that Mme. Tardieu-Blot (1958) 
described as A. quezelii. It came from Mt. Emi-Koussi in the Tibesti massif of northern 
Chad (Sahara desert). She compared her new species only with A. lepidum Presl, a 
distantly related south European fern. A. quezelii was also mentioned by Reichstein et 
al. (1 973), and compared with A. creticum. Most later authors (Quezel (1 971 : 448 & 
1983 : 414), Ozenda (1977 : 524)) have followed Tardieu-Blot's species concept (= an 
endemic of the Tibesti plateau, related to A. lepidum). 

Since A. daghestanicum and A. quezelii look very similar, a comparative 
morphological study, including macro- and micro-characters, was undertaken to 
establish whether they were closely related or conspecific. 

MATERIALS AND METHODS 
The following collections were studied: 
A. daghestanicum - Alexcenko et Woronow 450, U.S.S.R., Daghestan, Kurinski 

District, near Kuraeg, 21-V-1902. (Holotype P !, iso BR !). 
A. daghestanicum - Askerov s.n., U.S.S.R., Daghestan, 2200m. 1982 (LE !). 
A. daghestanicum - Reichstein 6062 (TR, 6062), cultivated progeny of Askerov s.n., 

Basel (GENT, pers. herb. T. Reichstein). 
A. quezelii - Quezel s.n., Chad, Tibesti, Emi-Koussi, lappiaz volcaniques du 

flanc NW, 3300m, 1956. (Holotype P !). 



144 



FERN GAZETTE: VOLUME 13 PART 3 (1987) 




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ASPLENIUM QUEZELII IDENTICAL WITH A. DAGHESTANICUM 



145 




FIGURE 2. Paleae.A daghestanicum [A\excenko et Woronow450; (a : holo-,b: isotype); Askerov 
s.n. (c,d,f-h); TR 6062 (e)], A. quezelii [Quezel s.n.-holotype (i-m)]. Bar = 1 mm. 



146 



FERN GAZETTE: VOLUME 13 PART 3 (1987) 



Epidermis preparations were made and studied following standard procedures 
(Viane 1985); SEM investigations were done on untreated spores coated with gold. All 
micromorphological characters used are mentioned in Table 2. Stomatal terminology 
is according to Van Cotthem (1970). 

RESULTS 
Macromorphological analysis 

The overall habit of the plants studied is the same (Fig. 1 ): all have a short oblique to 
upright rhizome, with several frondsclose together atthe top. The paleae(rhizome and 
stipe base) of both species are identical (Fig. 2); clathrate, without a central dark line, 
and with a few marginal outgrowths, there are no large size differences (1 .5-3mm long 
x 0.4mm wide) between the specimens. The leaves (3-7cm long) have a stipe about 
twice as long as the lamina (1 .5-2. 5cm long x 1cm wide), which isbipinnateonlyatthe 
base. The pinnae, gradually reduced towards the confluent apical segment, are a little 
longer than broad; their dorsal surface is often obscured with sporangia. The indusia 
(0.8-1 .5mm x 0.4mm) have an entire to slightly undulating edge. Table 1 gives the 
mean values and ratios for the macroscopical characters studied; all collections 
closely agree. The only discordant element is the short stipe of thecultivatedplant(TR- 
6062): the reduction of the stipe length is an apparently very common effect of 
cultivation (compare values from the original, wild (Askerov) collection with these of 
TR 6062 (its progeny)). 

Micromorphological analysis 

The pattern of the epidermis cells is so similar for all collections that only that of the 
holotypes is illustrated (Figure 3). Stomates, of the polocytic type, haveguardcells40- 
50um long. The polocytic cell, this is the cell surrounding the guard cells distally, is 
always a little wider than long. The exospore length is 30-36jjm, which indicates, just 
as the guard cell size, that the specimens are probably tetraploid. All microcharacters 
studied are on Table 2; it is clear that none of the col lections can be separated from the 
set on microscopical grounds. 




w^\- 




FIGURE 3. Epidermal cell patterns : A .A. daghestanicum (holotype); B : A. quezelii (holotype). Bar 
= 100jjm; arrow indicates direction of vein towards leaf margin. 



ASPLENIUM QUEZELII IDENTICAL WITH A. DAGHESTANICUM 



147 



Scanning electron microscopy (SEM) of spores can often be used to distinguish 
closely related taxa (e.q.Aspfenium adiantum-nigrum L from A cuneifolium Viv. and 
A onopteris L; A. fontanum ssp. fontanum from/1, fontanum ssp. pseudofontanum 
(Koss.) Reichst. & Schneller; etc.). SEM of A. daghestanicum and A quezelii spores 
(Fig. 4) revealed that these are so similar that they cannot be used to separate the 
species, not even on a subspecific or varietal level. 




FIGURE 4. SEM spore picture of A. daghestanicum (A); and/4, quezelii (B). Bar = 10pm. 

Conclusion 

The conclusion of this morphological study is that/4, daghestanicum and/4, quezelii 
are the same species. The correct name must then be A. daghestanicum, as Christ's 
binomial is the oldest legimate one; A. quezelii Tardieu-Blot is hereby reduced to a 
(taxonomical) synonym. 

DISTRIBUTION AND RELATIONSHIPS 
Until now, both A. daghestanicum and A. quezelii were considered endemics, the 
former of Daghesfan and the latter of the Tibesti massif. Following Tardieu-Blot's 
(1958) description and statements about the supposed affinities of A. quezelii with the 
south european A. lepidum, most later authors have copied her in their 
phytogeographic papers [a.o. Knapp (1 973 : 425); Ozenda (1 977 : 523); Quezel (1 971 : 
448, 1 983 : 414) etc.]. In a critical article Lebrun (1 983) showed that the number of true 
endemic species (12) from the Saharan mountains is considerably lower than had 
been estimated (85). Now A. quezelii can also be added to his list of so-called pseudo- 
species (= pseudo-endemic species). 

The former considerable phytogeographic interest of A. quezelii is not lessened 
now that it is included in A. daghestanicum. At present it is only possible to consider 
the African population as a Caucasian element in the Tibesti flora; it(both) probably is 
(are) relic(s)of a flora that was present under much wetter conditions. The author does 
not know of any other plant with a similar disjunct (3800km) distribution. 

A. daghestanicum is related to a number of (partly undescribed) small ferns, best 
represented in the Himalayas and China (a.o. A. kongashanense Ching, A. 
subdigitatum Ching, A. xinjangense Ching). Some new species belonging here will be 
described in the near future (Reichstein et al. in press). In the meantime, plant 
collectors in the Mediterranean to S.E.Asia are asked to look out for these rather small, 
easily overlooked ferns. 



148 



FERN GAZETTE: VOLUME 13 PART 3 (1987) 





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ASPLENIUM QUEZELII IDENTICAL WITH A. DAGHESTANICUM 149 



ACKNOWLEDGEMENTS 
I thank the Directors and Curators of BR and P for the loan of valuable type material. 
Prof. Reichstein supported and encouraged my work in many ways; he also cultivated 
and provided material of the Askerov collection. Ir. Bohyn made the SEM available. I 
thank Prof. Van Cotthem and Prof. Van der Veken for supporting my research at their 
laboratory. I am grateful to Dr. Mary Gibbyfororganizingthisspecial anniversary issue 
and for revising the English text. 

REFERENCES 

CHRIST, H. 1906. Deux fougeres nouvelles du Caucase. Mon. Jard. Bot. Tiflis 10: 24-25. 

KNAPP, R. 1973. Die Vegetation von Afrika. Fischer Verlag, Stuttgart. 

LEBRUN, J. -P. 1 983. La flore des massifs Sahariens, especes illusoires et endemiques vraies. 

Bothalia 14: 511-515. 
OZENDA, P. 1 977. Flore du Sahara (2nded.). Centre National de la Recherche Scientifique, Paris. 
OUEZEL, P. 1971. Flora und Vegetation der Sahara. In H. Schiffers (Ed.), Die Sahara und ihre 

Randgebiete, band I. Physiogeographie. Weltforum Verlag, Munchen. 
QUEZEL, P. 1983. Flore et vegetation actuellesde I'Afriquedu Nord, leur signification en fonction 

de I'origine, de revolution et des migrations des flores et structures de vegetation 

passees. Bothalia 14: 41 1 -41 6. 
REICHSTEIN, T., LOVIS, J.D., GREUTER, W. & ZAFFRAN, J. 1 973. Die Asplenien der Insel Kreta. 

Ann. Mus. Goulandris 1: 133-163. 
TARDIEU-BLOT, M.-L. 1958. Etudes des Pteridophytes du Tibesti et du Borkou. In P. Quezel, 

Mission botanique au Tibesti. Mem. Inst. Recherches Sahar. 4: 90-91 . 
VAN COTTHEM, W. 1970. Comparative morphological study of the stomata in the Filicopsida. 

Bull. Jard. Bot. Nat. Belg. 40: 81 -239. 
VIANE, R. 1985. Dryopteris expansa and D. x ambroseae (Pteridophyta) new for Belgium. Bull. 

Soc. Roy. Bot. Belg. 118: 57-67. 



15 o FERN GAZETTE: VOLUME 13 PART 3 (1987) 



REVIEW 

FERNS AND ALLIED PLANTS OF VICTORIA, TASMANIA AND SOUTH 
AUSTRALIA by Betty D. Duncan and Go/da Isaac. 258+xiipp., 246 figures, 92 
maps, 8 plates, 185 x 255mm. Melbourne University Press. ISBN: 522 
84262 3. Price £25.00. 

This book is a welcome addition to the ranks of fern floras and certainly lives up to the 
claim on its dust cover to be of use to naturalists, botanists and gardeners. It is 
intended to be used primarily as an aid inthe field identification of ferns andfern allies. 
Two keys to genera are provided. The first is a tabular key for fertile material which is 
conveniently printed on a single fold out page. The second is an illustrated 
dichotomous key for fertile and sterile specimens. Within each genus there is a key to 
species and these keys, together with detailed descriptions of each species, clearly 
indicated field characters and good black and white photographs or drawings of whole 
fronds or complete plants (and sometimes more detailed illustrations of rhizomes, 
scales and fertile parts of the frond) should enable anyone to make correct 
identifications. The colour plates provide additional important information on habit, 
habitat and frond colouration; they are of the same high quality we have come to 
expect from recent Australian fern books. Distribution maps are included for most of 
the species which occur in the state of Victoria and much useful ecological information 
is provided. For the gardener there are guidance notes on situation requirements and 
suitability for pot, basket or rockery cultivation, while the broader principles, both of 
propagation by vegetative means and by spores, and of fern growing in general, are 
dealt with in the final chapter by C.J. Goudey and R.L. Hill. 

The authors are to be congratulated on their high standards of content and 
presentation. I highly recommend this book to anyone with an interest in Australian 
pteridophytes and indeed to any naturalist intending to visit south-eastern Australia. 
At f 25.00 it is very reasonably priced. 

B.S. PARRIS 



FERN GAZ. 13(3) 1987 151 



HYBRIDISATION EXPERIMENTS BETWEEN 

ASPLENIUM SEELOSII AND A. CELTIBERICUM 

(= A. SEELOSII SUBS P. G LAB RUM) 

(ASPLENIACEAE: PTERIDOPHYTA) 



J.D. LOVIS 

Department of Plant and Microbial Sciences, University of Canterbury, 

Christchurch, New Zealand* 

ABSTRACT 
Hybridisation experiments between Asplenium seelosii Leybold from the Italian 
Dolomites and/4, celtibericum Rivas- Martinez {-A. seelosii subsp. glabrum (Litard. 
& Maire) Rothm.) from N.E. Spain are described. The F, hybrids showed a regular 
meiosis (n = 36) and were fertile. F 2 generations were raised from two of the F : 
plants. These showed indications of recombination of factors affecting fitness. The 
decision by Reichstein to revert to subspecific status for the Iberian taxon is upheld. 

Asplenium seelosii sensu lato is one of the strangest and most distinctive elements in 
the very diverse European spleenwort flora (cf. Eberle 1959, p. 66; Rasbachetal. 1968, 
fig. 48, p. 101). The only species to which it has even a remote resemblance is A 
septentrionale (L)Hoffm., with which it shares the feature of a highly reduced lamina. 
Indeed, it was initially regarded as a variety (var. tripartitum) of that species by Gustav 
Seelos, after whom the plant was subsequently named when described as a distinct 
species by Leybold (1855). (See Becherer 1962, p. 55). 

Although so distinctive, Asple nium seelosii is nevertheless highly variable, not 
merely between and within populations, but also within one individual, fronds varying 
in morphology in response to environmental conditions, including climatic variation 
and season of the year. This variability is not suppressed by glasshouse culture. In 
these experiments, plants grew more luxuriantly in Basel than in Leeds, though the 
growth forms produced in Leeds were perhaps closer to those likely to be realised in 
the wild. 

In its more luxuriant forms, the fronds oi Asplenium seelosii consist of a relatively 
long stipe, terminated by a very short lamina divided into three segments, which may 
themselves be incised to a greater or lesser extent. The frond is rarely quite 
symmetrical, one of the lateral segments usually joining the axis at an appreciably 
lower point than the other. 

The characteristic habitat of Asplenium seelosii is crevices of vertical limestone 
rock walls. Its distribution falls into two main highly disjunct areas. In central Europe, 
its range is based on the Italian Dolomites with outliers in Austria, N.W. Jugoslavia 
(Julian Alps) and Germany (just!). It occurs also in the N.E. quarter of Spain, where 
there are two centres — one in the eastern half of the F\renees (one locality here is 
actually in France) and the other south of the Ebro river in Guadalajara. The plant is not 
endemic to Europe; it occurs also in -Morocco (cf. Jalas & Suominen 1972, p. 77). 

The type of Asplenium seelosii Leybold is from the Tyrol. Unless depauperate, 
plants from the Dolomites (Figs. 1A-I)always have a tripartite lamina, and this lamina 
plainly bears numerous stalked glandular hairs. In contrast, plants from Spain are 
glabrous and have a less divided lamina, though it is now evident that populations vary 
in this latter character (Figs. 1 J-S). These differences have resulted in Iberian plants 
being given separate taxonomic status, firstly as a variety, var. glabrum Litardiere & 
Maire (in Maire 1928), secondly as a subspecies, subsp. glabrum, by Rothmaler (in 

formerly of Department of Plant Sciences, University of Leeds. 



152 



FERN GAZETTE: VOLUME 13 PART 3 (1987) 




FIGURE 1. Silhouettes, * 0.875, of pressed fronds of Asplenium see/osii (A-\) and A. celtibericum 
(J-S). A-C) Salurn, Italian Dolomites, wild collection. D-F) Buco de Vela, Italian Dolomites, cult. 
Leeds. G-l) Mazzin, Italian Dolomites, wild collection. J-L) progeny of isotype of A. celtibericum, 
Pinar de Campisabalos, Guadalajara, Spain, legit Rivas-Martinez et al., cult. Leeds. M & N)TR 
898, N. of Organa, valley of R. Segre, N.E. Spain. O & P)TR 898, cult. Basel. Q-S)TR 1 193, chapel 
of Sta. Fee, near Organa, legit A. & C. Nieschalk, cult. Basel. 



HYBRIDISATION EXPERIMENTS BETWEEN ASPLENIUM SEELOSII & A. CELTIBERICUM 153 



Cadevall & Font Quer 1937) and lastly raised to full specific rank as A celtibericum by 
Rivas-Martinez (1967). A. celtibericum is based on material of extreme morphology 
from Guadalajara. Progeny of isotype material raised in Leeds is diminutive, with a 
virtually undivided lamina, consisting of only a single serrated terminal segment (Fig. 
1J-L). 

Both taxa are, as far as is yet known, uniformly diploid (Meyer 1 957, 1 967; Lovis 
unpub.). 

It is clearly a matter of some interest to establish the degree of genetical 
relationship persisting between these two taxa — their origin from a common ancestor 
surely cannot be doubted. Accordingly, two attempts were made to hybridise them, 
using a characteristic though not extreme form of the Spanish plant as the male 
parent. The overall success obtained was 50%. 



FEMALE PARENT 



MALE PARENT 



FEMALE 
PROTHALLI 



HYBRIDS 



JDL 1439 



A. seelosii Mazzin, 
Italy 



A. celtibericum Organa, 

Spain (TR 898) 



10 



JDL 1440 



10 



20 



10 



The Ft hybrids were unexpectedly easy to discriminate from possible 'female'selfs 
(or stray selfed 'male' prothalli), being not only vigorous, but of distinctive morphology. 
They all retained the tripartite lamina of Asplenium seelosii sensu stricto, but were 
glabrous. Thus in the F, hybrids division of the lamina was dominant, but glandulosity 
proved to be recessive. Chromosome pairing was regular, 36 bivalents being formed, 
as in the two parents. F 2 generations were raised from two of the F } hybrids. In contrast 
to the uniform morphology of the F : hybrids (Figs. 2A-E, 3A-F), the F 2 progenies (Fig. 
3G-X) were variable in frond form, with some individual recombinants lying outside 




FIGURE 2 Silhouettes, * 0.875, of pressed fronds of five synthetic hybrids, A. seelosii * 
celtibericum (F,), all cult. Leeds. A) JDL 1439A. B) JDL 1439D. C) JDL 1439B. D) JDL 1440B. 
E) JDL 1440C. 



15 4 FERN GAZETTE: VOLUME 13 PART 3 (1987) 



the range of the parent taxa (e.g. Fig. 3M-P). Curiously, only the sparest indications of 
glandular hairs were seen on the lamina of any of the F 2 plants. The suppression of 
glandulosity in both hybrid generations suggests that this characteristic evolved in the 
Tyrolean plant subsequent to its separation and divergence from the Iberian taxon and 
was not present in their common ancestor. The F 2 progenies also showed signs of 
recombination of factors affecting fitness, numerous sporophytes not surviving to 
maturity. Success in raising in the glasshouse a plant from so extremely specialised a 
habitat as that of A seelosii is at best precarious, but nevertheless there is good reason 
to believe that, in practice, the F 2 generation shows some reduction in vigour in 
comparison with the F^ The fertility of F 2 plants was not tested. 

In the light of this information, combined, I suspect, with his personal experience 
of the range of form present in Spanish populations, Reichstein, in his remarkable 
definitive review of hybrids in European Aspleniaceae (Reichstein 1981, p. 105), opted 
for subspecific status for the Iberian plant. Taxonomic decisions of this type can be 
more a matter of taste, of personal preference, rather than a question of right or wrong, 
but nevertheless Reichstein's decision is plainly very appropriate. Although it is true 
that the two areas of distribution of these taxa are so disjunct that their interfertility 
can have no practical effect in nature, recognition of the two taxa as geographical 
subspecies has the clear merit of preserving, within the classification, an indication of 
the close relationship that undoubtedly exists. 

A remarkable diversity of form exists within the diploid taxa present in European 
Aspleniaceae. Using the ability to pair chromosomes at meiosis as a criterion of 
relationship, the great majority of inter-specific diploid combinations so far studied 
show no evidence of affinity. This can be true (e.g. tr/chomanes xviride : 72 univalents) 
or virtually so (e.g. cuneifolium & onopteris : - 6 bivalents*), even where some 
evidence of affinity might be expected on morphological grounds. In contrast, A. 
seelosii x celtibericum shows us a situation where geographical isolation exists, 
accompanied by some morphological differentiation, but divergence is not yet 
sufficient to affect chromosome pairing. 

It would be of great interest to investigate certain other pairs of diploid species of 
Asplenium, namely \)A.jahandiezii (Litard.) Rouy and/4, bourgaei Boiss. ex Milde,and 
2) A. aegaeum Lovis, Reichst. & Greuter & A fissum Kit. ex Willd. A. jahandiezii is 
restricted to the vicinity of the Gorge du Verdon in S.E. France, whereas A bourgaei is 
a plant of Asia Minor, principally Asian Turkey. Their morphology suggests a close 
relationship, though by no means as close as in the case of A. seelosii and A. 
celtibericum. Although A. aegaeum and A. fissum are unmistakeable in their most 
characteristic forms, a few collections are difficult to place. Their distributions are 
approximately contiguous in the Aegaean region. All of these four species are difficult 
to cultivate, and it has not yet been possible to attempt successfully to hybridise them, 
using the* technique employed in Leeds (Lovis 1968), for which a prothallial growth 
sufficiently dense and vital to show, when tested, some signs of spermatozoid 
liberation is necessary if an hybridisation attempt is to have a reasonable chance of 
success. For the synthesis of such hybrid combinations, the elegant 'nearest 
neighbours' transplant technique devised and exploited with success in recent years 
by Reichstein, wherein the opportunity for hybridisation isextended indefinitely rather 
than restricted to a brief span in a watchglass, may well offer much better prospects. 



*Evidence from X Asplenophyllitis jacksonii Alston (Lovis &Vida 1969), A xbechereriD.E. Meyer 
and A xdolosum Mi Ide (Reichstein 1981 , Appendix II, pp. 1 18 & 122). Note that tetraploid hybrids 
such as A major/cum * adulterinum which show 144 univalents (Lovis & Reichstein 1969) in 
effect provide a demonstration of lack of ability to pair for no less than six different diploid 
combinations : PF, PT, PV, FT, FV & TV. 



HYBRIDISATION EXPERIMENTS BETWEEN ASPLENIUM SEELOSII & A. CELTIBERICUM 155 




u 



w 



FIGURE 3 Silhouettes, x 0.875, of pressed fronds of two synthetic hybrids, Asplenium seelosu * 
celtibericum (A-F), and their selfed F 2 progeny (G-X). A-C) JDL 1439C, cult. Leeds. D-F) JDL 
1 439E (TR 1 524), cult. Basel. G-P) selfed progeny of JDL 1 439C, cult. Leeds. G & H) JDL 1 439C:1 . 
I&J)JDL1439C:2. K) JDL 1439C:3. L)JDL 1439C:6. M-P) JDL 1439C:s.n. Q-X) selfed progeny of 
JDL 1439E, cult. Leeds. Q-S) JDL 1439E:1. T & U) JDL 1439E:2. V-X) JDL 1439E:s.n. 



156 



FERN GAZETTE: VOLUME 13 PART 3 (1987) 



ACKNOWLEDGEMENT 
It is a great pleasure and privilege to acknowledge the assistance of Prof. Dr. Tadrik 
Reichstein in this project, 1) in guiding me to localities of A. seelosii s.s., including that 
at Mazzin from whence the culture used as female parent was obtained, 2) for 
providing the material of subsp. glabrum subsequently utilised as male parent and 
3) for cultivating in Basel, very successfully, one of the resultant Ft hybrids. 

The great increase in our knowledge of European Aspleniaceae in the last 25 years 
is in very large measure the achievement of one man, both directly by his own efforts, 
and otherwise by very materially assisting and inspiring others. This paper is dedicated 
to him with most grateful thanks by one who has been very fortunate in being an 
associate. 



REFERENCES 

BECHERER, A. 1962. Uber die geographische Verbreitung von Asplenium seelosii Leybold. 

Bauhinia 2: 55-58. 
CADEVALL, J. & FONT QUER, P. 1936. Flora de Cata/unya 6. Barcelona. 
EBERLE, G. 1959. Fame im Herzen Europas. Frankfurt. 

JALAS, J. & SUOMINEN, J. 1972. Pteridophyta. Atlas Florae Europaeae 1. Helsinki. 
LEYBOLD, F. 1855. Asplenium seelosii, ein neuer Farm aus Sudtirol. Flora 38: 81-82. 
LOVIS, J.D. 1968. Fern hybridists and fern hybridising. II. Fern hybridising at the University of 

Leeds. Brit. Fern Gaz. 10: 1-8. 
LOVIS, J.D. & REICHSTEIN, T 1969. Der Fambastard Asplenium x orel/ii hybr. nov. = Asplenium 

majoricum Litard. x A. trichomanes L. subsp. quadrivalens D.E. Meyer und die 

Abstammung von A. majoricum. Ber. Schweiz. Bot. Ges. 79: 335-345. 
LOVIS, J.D. & VIDA, G. 1 969. The resynthesis and cytogenetic investigation oiXAsplenophyllitis 

microdon and X A. jacksonii. Brit. Fern Gaz. 10: 53-67. 
MAIRE, R. 1928. Contributions a I'etude de la flore de I'Afriquedu Nord. Bull. Soc. Sc. Nat. Maroc 

8: 128-143. 
MEYER, D.E. 1957. Zur Zytologie der Asplenien Mitteleuropas (\-XV).Ber. Deutsch. Bot. Ges. 70: 

57-66. 
MEYER, D.E. 1 967. Uber neue und seltene Asplenien Mitteleuropas, 4. Mitteilung.fier. Deutsch. 

Bot. Ges. 80: 28-39. 
RASBACH, K., RASBACH, H. & WILLMANNS, O. 1968. Die Farnpf/anzen Zentra/europas. 

Heidelberg. • 
REICHSTEIN, T. 1981. Hybrids in European Aspleniaceae (Pteridophyta) Bot Helvetica 91: 89- 

139. 
RIVAS-MARTINEZ, S. 1967. Une espece nouvelle dAsplenium (Aspleniaceae) d'Espagne. Bull. 

Jard. Bot. Nat. Belg. 37: 329-334. 



FERN GAZ. 13(3) 1987 157 



OBSERVATIONS OF PROGENY OF ATHYRIUM FILIX-FEMINA 
(ATHYRIACEAE; PTERIDOPHYTA) 
FROM BREEDING EXPERIMENTS* 



J.J. SCHNELLER 

Institut fur Systematische Botanik, Zollikerstr. 107, CH-8008 

Zurich, Switzerland 

ABSTRACT 

Progeny of plants collected from natural Swiss populations have been grown for 
more than 8 years in a garden. The offspring resulting from intragametophytic 
selfing are on average much smaller and more variable in size than those from 
intergametophytic selfing or from crossing (out-breeding). The sporophytes 
originating from outbreeding were the tallestand most robust. This observation and 
earlier results (Schneller 1 979) suggest that the genetic variability and genetic load 
may be very high. How is it possible to maintain so many recessive deleterious 
genes in natural populations? Because of the instability of the microhabitats in 
which the prothalli grow, soft selection may allow an accumulation of genetic load. 
The result of this investigation, however, can also be interpreted in light of genetic 
regulation, because leaf dimensions are normally controlled by polygenic 
complexes. 

INTRODUCTION 
In most homosporous ferns it is possible to breed sporophytes in three different ways: 
1. intragametophytic selfing, 2. intergametophytic selfing, 3. intergametophytic 
crossing. In the first case completely homozygous sporophytes result, because the egg 
and spermatozoid are from the same prothallus and therefore genetically identical. 
Intergametophytic selfing implies mating gametes from two prothalli of the same 
parent plant. As a result totally homozygous or partially heterozygous progeny will 
originate depending on the mode of fertilization and the genetic composition of the 
parent plant. In intergametophytic crossing experiments prothalli from two different 
parent plants are used. In this case either homozygous plants (due to 
intragametophytic selfing) or heterozygous plants (degree of heterozygosity depending 
on the genetic constitution of the parents) result. In a study of the breeding system of 
Athyrium filix-femina (Schneller 1979) it was shown that outbreeding was most 
successful but intra- and intergametophytic selfing also led to progeny. The viability of 
the different kinds of progeny, however, was remarkably different. After 8-10 years 
some of the plants obtained in these earlier experiments are still alive and are growing 
under similar conditions. Progeny from the three different modes could be seen to be 
distinct. The reasons for the observed differences will be discussed. 

MATERIALS AND METHODS 
The parent plants were collected in Switzerland in natural populations at 
Horgenerberg, Horgen, Kt. Zurich (nr. A-1 , A-2, A-3, A-29, A-31, A-33), at Scalasiten, 
Tamins, Kt. Grisons (nr. A-42, A-48, A-49, A-50), and at Goscheneralp, Goschenen, 
Kt. Uri (nr. A-51). The plants investigated here were experimentally produced in the 
years 1 975-77. Since 1 979 they have been grown under similar conditions in a shady 
garden bed in the Botanical Garden of the University of Zurich. Three classes of plants 
were evaluated: 1. progeny (18 plants) from intragametophytic selfing ('intra'), 2. 
progeny (21 plants) from intergametophytic selfing experiments ('inter'), and 3. 
progeny (12 plants) from crossing experiments ("cross"). In an earlier investigation 

*Dedicated to Prof. Dr. T. Reichstein en his 90th birthday. 



158 



FERN GAZETTE: VOLUME 13 PART 3 (1987) 



intra 






















• 


• 




• 




o 


• 


• 


• 


• 




• 


• 


• 



50 



100 cm 



inter 





I — 


— h 




H 






o 


o 








o 


• 










• 


• 






o 


o 


• 


• 




• 

















50 



100 cm 



cross 



i — i — i 





o 








• 






• 








• 


o 






® 


• 




o 


® 


® 


® 



50 



100 



cm 



FIGURE 1 . Distribution of leaf lengths of the different types of progeny, intra = intragametophytic 
self ing, inter = intergametophytic self ing, cross = crossing, o = progeny from plant A- 1 , • = progeny 
from plant A-2, = progeny resulting from crossing A-1xA-2, ,_,_,= mean and standard deviation, 
blanks = progeny from plants other than A1, A2. 



OBSERVATIONS OF PROGENY OF ATHYRIUM FILIX-FEMINA 159 



(Schneller & Schmid 1 982) and again in this experiment it could be shown that the 
aspects of leaf size that determine shape are correlated. This explains why the size of 
the leaves was used for Figure 1. Methods for the statistical analysis of means and 
deviation tests follow Under & Bechtold (1 979) and Campbell (1971), SAS statistical 
programs (SAS Inst. 1982) were used. 

RESULTS 
Measurements given in Table 1 were used for the statistical comparison. The Kruskall- 
Wallis test showed significant differences between 'intra' and 'cross' and between 
'inter' and 'cross' (SAS Inst. 1982) In Figure 1 the distribution of the leaf length is 
shown. The progeny of two plants (A-1 and A-2) are indicated. Principal component 
analysis shows significant differences between the different breeding-classes. 

DISCUSSION 
Earlier experiments (Schneller 1979) showed that Athyrium filix-femina is 
predominantly outbreeding. All the parent plants collected randomly from natural 
habitats and used for the breeding experiments were heterozygous. These original 
plants grew well in their original environment. They can be regarded as successful 
results of selection (in the past) because they survived and withstood competition. 
Earlier experiments (Schneller 1979) and the results presented here show that the 
original plants stored a remarkable amount of genetic variability. In this investigation 
(Fig. 1 ) it was shown mainly in examples A-1 and A-2 (but is true also for other 
examples). Because progeny are growing under similar conditions much of the 
observed variability is due to genetic diversity (it is difficult or impossible to distinguish 
in nature between the genetic component of variation and phenotypic plasticity 
(Schneller & Schmid 1982) because they are intermingled). Genetic variability maybe 
expressed by advantageous, neutral, or deleterious factors. The results of breeding 
experiments reveal some consequences of this genetic diversity. Intragametophytic 
and to a lesser degree intergametophytic selfing show very different results. In many 
cases no sporophytes are formed. If sporophytes occur they differ strikingly. They may 
die at an early stage or vary in size and virility (Schneller 1979). When no 
sporophytes are observed this may be due to prezygotic mechanisms or to early lethal 
recessive genes (zygotic iethals). Dwarf or subvital sporophytes may be due to weaker 
recessive deleterious genes. 

After nearly ten years it can be clearly seen that the three classes of progeny 
('intra', 'inter' and 'cross'; Fig. 1) are distinct considering standard deviation and 
means of length. The differences are statistically significant with the exception of the 
mean leaf-length of 'intra' and 'inter' (which is significant only at the 80% level). The 
class 'intra' shows the largest deviation and the greatest proportion of dwarf plants. It 
should be noted that intragametophytic selfings have the greatest number of deaths 
during early stages (Schneller 1979). From the 50 original "intra"-plants only 18 
survived. Even the progeny of one plant (A 1)showsa wide rangeof different offspring. 
Because the plants are homozygous, recessive genes will always be expressed. The 
great variance reflects genetic diversity. Dwarf or small plants couldthen bethe result 
of weakly deleterious recessive genes, that may influence the metabolism of these 
plants. The viability of the plant and the regulation of leaf size seem to be somewhat 
correlated. 

In the class 'inter' the deviation is smaller and the mean is larger than in the class 
'intra'. This would support the hypothesis of increased heterozygosity. Heterozygosity 
in this case seems to have a stabilising effect on the development, i.e., some of the 
recessive deleterious alleles will be "masked" by dominant alleles. Some of the 



160 FERN GAZETTE: VOLUME 1 3 PART 3 (1 987) 



recessive deleterious genes, however, will be found in homozygous condition. This 
would explain the still wide range of different phenotypes in this class. We can assume 
that the degree of heterozygosity is highest in the class 'cross'. Here we observe taller 
plants, a smaller variability and a greater viability. The difference between the other 
two origins can be explained again by the stabilising effect of heterozygosity. 

In population genetics simplified models have often been used to explain the 
influence of genes on fitness (Sperlich 1973, Wallace 1970). The results of this 
investigation in my opinion cannot be explained satisfactorily by simple models. 

It is obvious that genetic load (mutational and/or segregational) occurs in the 
gene pool of Athyrium filix-femina. It has to be regarded as a consequence of 
outbreeding. Thus, stored recessive genetic load will be a hindrance to inbreeding, 
mainly because lethal genes result in none or in subvital offspring that could not 
survive in natural conditions. Most inbred sporophytes could not compete with the 
crossbred ones. Fern geneticists like Lloyd (1974) and Klekowski (1982) argue that 
inbreeding is important for the establishment of new -populations. Selection pressure 
for inbreeding would 'clean' the population of deleterious genes and would also have 
an influence on the genetic diversity. The genetic load observed in this example 
reflects a great genetic diversity. Is such a genetic load extraordinary for ferns (or other 
organisms)? We still do not know enough abouthow much of genetic diversity can be 
maintained in natural populations. The answers given by theoretical models range 
widely depending on the assumptions made. Looking at concrete results in ferns, 
Klekowski (1982) mentioned that Osmunda regalis has the highest load so far 
documented. For this species 2.39 lethal equivalents per zygote were calculated. A 
somewhat lower load was found in Thelypteris palustris (Ganders 1972). Based on my 
earlier results (Schneller 1979) and resultsof this investigation lestimate (method see 
Ganders 1972) a mean. load of 4.87 lethal equivalents per zygote for Athyrium filix- 
femina. This is about double the load of Osmunda regalis (Klekowski 1982). A 
comparable amount of load can be found in human populations (Brues 1969). 

Wallace (1970, 1975) showed that soft selection (which is density- and 
frequency-dependent) allows an accumulation of genetic diversity. The consequences 
of soft selection to genetic diversity and particularly to genetic load were calculated by 
Klekowski (1982). In my opinion it is the habitat of A. filix-femina (and of course other 
plants as well) that allows soft selection. It consists of a mosaic of microhabitats 
variable within some limits influenced by other organisms or by unpredictable physical 
events on the soil, such as erosion, disturbance by rain, snow, frost, cover of litter etc. 
The observed large genetic diversity may be partly duetothe natureof the microhabitat 
and soft selection. 

The conclusions or hypotheses made so far in this discussion were based on the 
assumption that the observed load is due to recessive lethal genes. We know for 
numerous plants that the regulation of size and shape is polygenic (Mather 1 942). The 
different leaf sizes seen in the experiments (Fig. 1) could also be explained by the 
different combination of regulating genes and could be a consequence of epistasis. We 
still know little about either the interrelationship of structural and regulatory factors or 
the role of epistasis. 

Finally, I am fully aware of the lack of a satisfactory explanation to the observed 
phenomena. But in my opinion we should focus our interest on the problems of gene 
regulation; some attributes we currently link with genetic load ma/ have another 
cause. 

ACKNOWLEDGEMENTS 
I thank Larry Hufford for reading and correcting the manuscript, Mark Nicholls and 
Prof. C.D.K. Cook for comments. 



OBSERVATIONS OF PROGENY OF ATHYRIUM FILIX-FEMINA 



161 



TABLE 1 



intra 


inter 


cross 


Nr. 


L 


B 


Ba 


Pos 


Nr. 


L 


B 


Ba 


Pos 


Nr. 


L 


B 


Ba 


Pos 


A1h1 


104 


14.0 


7.5 


10 


A1M 


76 


12.5 


5.5 


7 


A1/A2-2 


106 


17.0 


9.0 


7 


A1h3 


78 


12.0 


5.5 


8 


A 1i4 


60 


9.0 


5.0 


8 


A 1 /A 2-3 


92 


14.5 


7.5 


9 


A1h4 


71 


10.5 


5.0 


8 


A1i5 


71 


11.0 


4.0 


6 


A1/A2-4 


91 


15.0 


6.0 


6 


A1h5 


12 


1.5 


1.0 


4 


A1i7 


71 


10.5 


5.0 


8 


A1/A2-5 


88 


13.0 


5.5 


7 


A1h6 


78 


11.5 


5.5 


7 


A1i8 


81 


12.0 


4.5 


7 


A1/A31-1 


89 


13.5 


5.5 


9 


A1h7 


67 


11.0 


6.0 


7 


A1i12 


99 


13.5 


5.5 


7 


A1/A31-4 


91 


15.0 


9.0 


6 


A1h8 


42 


6.5 


4.5 


8 


A1M3 


83 


11.5 


5.5 


5 


A1/A42-2 


94 


16.0 


7.0 


8 


A1h9 


58 


9.5 


4.0 


6 


A1I14 


70 


11.0 


3.0 


8 


A1/A43-5 


93 


15.5 


5.5 


10 


A1h10 


38 


5.5 


2.5 


7 


A1I17 


92 


13.0 


9.0 


7 


A2/A31-1 


98 


14.5 


5.0 


9 


A1M1 


71 


11.5 


5.5 


8 


A1i19 


49 


8.0 


3.5 


8 


A2/A31-3 


92 


14.5 


7.5 


9 


A1h12 


84 


10.0 


6.0 


9 


A1I21 


104 


14.0 


6.5 


9 


A2/A48-1 


77 


13.0 


4.0 


7 


A2h1 


60 


8.5 


3.0 


6 


A1122 


77 


13.0 


8.5 


6 


A48/A47-C 


65 


12.5 


5.0 


7 


A3h5 


75 


13.0 


5.0 


7 


A2i2 


69 


15.0 


7.0 


5 












A29h2 


58 


8.5 


2.5 


8 


A2i3 


79 


13.5 


4.5 


8 












A33h1 


109 


16.5 


10.5 


7 


A2i4 


74 


15.0 


4.5 


9 












A33h2 


73 


12.5 


6.5 


5 


A2i6 


64 


9.0 


4.0 


6 












A48h3 


46 


7.5 


3.5 


7 


A2110 


48 


8.5 


2.5 


9 












A50h1 


29 


4.0 


3.0 


5 


A31i1 
A31i2 

A50I4 
A51i8 


45 
83 
59 
89 


9.0 
16.0 
11.0 
15.0 


4.0 

6.0 

4.5 

11.0 


5 
6 
5 
4 













TABLE 1. Leaf dimensions of progeny from different breeding experiments, intra = 
intragametophytic selfing, inter = intergametophytic selfing, cross + crossing, Nr. = plant number, 
L = length of leaf, B = breadth of leaf, Ba = length of basal pinna, Pos = position of longest pinna 
(basal pinna = Pos 1). 



REFERENCES 
BRUES, A.M. 1969. Genetic load and its varieties. Science 164: 1130-1136. 
CAMPBELL, R.C. 1971. Statistische Methoden fur Biologie und Medizin. G. Thieme Verlag, 

Stuttgart. 
GANDERS, F.R. 1972. Heterozygosity for recessive lethals in homosporous fern populations: 

Thelypteris palustris and Onoclea sensibilis. Bot. J. Linn. Soc. 6b: 21 1-221. 
KLEKOWSKI, E.J. 1982. Genetic load and soft selection in ferns. Heredity 49: 191-197. 
LINDER, A. & BERCHTOLD, W. 1979. Elementare statistische Methoden, UTB 796. Birkhauser 

Verlag, Basel. 
LLOYD, R.M. 1974. Genetic load in pioneer and non-pioneer Hawaiian pteridophyta.v/ Linn. Soc. 

Bot. 69: 23-35. 
MATHER K 1942. Polygenic inheritance and natural selection. Biol. Rev. 18: 32-64. 
SAS Inst.' Inc. SAS User's Guide Statistics 1982. SAS Institute. Cary, North Carolina. 
SCHNELLER J J. 1979. Biosystematic investigations on the lady fern (Athyrium filix-femma). 

Plant Syst. Evol. 132: 255-277. 
SCHNELLER, J.J. and SCHMID, B.W. 1982. Investigations on the intraspecific variability in 

Athyrium fi/ix-femina (L.) Roth. Bull. Mus. natn. Hist, nat., Paris, 4eser., 4 sect. B: 215- 

228. 
SPERLICH, D. 1973. Popu/ationsgenetik. G. Fischer Verlag, Stuttgart. 
WALLACE, B. 1970. Genetic load, its biological and conceptual aspects. Prentice-Hall, 

Englewood, N. Jersey. 
WALLACE, B. 1975. Hard and soft selection revisited. Evolution 29: 465-473. 



162 FERN GAZETTE: VOLUME 13 PART 3 (1987) 



REVIEW 

ICONOGRAPHIA PALYNOLOGICA PTERIDOPHYTORUM ITALIAE. E. 
Ferrarini, F. Ciampolini, R.E.G. Pichi Sermolli, D. Marchetti. 1986. Webb/a 
40(1): 1-202. ISSN: 0083-7792. 

This work is a study of the spores of the 1 22 species and subspecies of Pteridophytes 
found in Italy. The text is in Italian. For each taxa, a detailed morphological description 
of the spore(s) is given (a helpful glossary of the terms used, illustrated with line- 
drawings, is provided), with the global and Italian distributions, habitat information 
and cytological data. Although the (untreated) spores of all the taxa are illustrated by 
scanning electron micrographs, the spore measurements given are taken from dry 
spores (i.e. observed in air, without any mounting medium) with a transmitted-light 
microscope. The accounts of many of the taxa have comments on such topics as their 
taxonomy, nomenclature, cytology or phytogeography. A comprehensive bibliography 
of some 350 references is given. The new combination Polypodium cambricum ssp. 
serrulatum (Sch. ex Archang.) Pic. Ser. is made. 

J.M. CAMUS 






FERN GAZ. 13(3) 1987 163 



ASPLENIUM TADEI (ASPLENIACEAE : PTERIDOPHYTA) 
A NEW SPECIES FROM TURKEY 

C.R. FRASER-JENKINS 
71 Abingdon Road, Oxford, 0X1 4PR, England 

and J.J. SCHNELLER 

Institut fur Systematische Botanik, Universitat Zurich, Zollikerstra. 107, 

CH-8008, Zurich, Switzerland 

ABSTRACT 

The discovery of a new Asplenium species, A tadei Fraser-Jenkins & Schneller, is 
reported from Antalya province, south Turkey. Its possible relationships are briefly 
discussed in the light of its tetraploid cytotype, and it is concluded that it is probably 
an allotetraploid species, either involving A. trichomanes-ramosum L. and some 
other diploid species, possibly A. bourgaei, Milde or that it is a member of the A 
exiguum Bedd., or A daghestanicum Christ aggregates. 

INTRODUCTION 
In June 1 978, while examining specimens from western Asia at Kew (K), in connection 
with his work on "Flora Iranica" (Fraser-Jenkins, Khullar & Reichstein, in prep.), the 
first-named author came across an unusual specimen of an Asplenium from Turkey, 
which was generally similar to A viride Huds., but with an unusual frond-apex where 
the segments had become fused together into a terminal blade with narrow side-lobes. 
The specimen was unlike any other Asplenium species from the European area (sens, 
lat.) with the possible exception of A daghestanicum Christ, a rare species from the 
eastern Caucasus, then known only fromthe type specimens in Paris(P), andwhich he 
had previously spent some time attempting to obtain, both in the Caucasus and by 
correspondence, in connection with study on that species and the closely similar, 
Sino-himalayan A. aitchisonii Fraser-Jenkins & Reichstein, by Professor T. Reichstein 
of Basel. However the Turkish specimen was noticeably longer, narrower and less 
deeply dissected than the type material of A daghestanicum, and had the sori crowded 
towards the base of the pinnae; subsequently new material of A daghestanicum 
brought to the BM from Dr A.N. Askerov of Baku, with the help of Prof. A. Takhtajan of 
Leningrad, showed that it was not the same species. The specimen concerned had 
been cited in the "Flora of Turkey" (Davis (ed.) 1965) under A bourgaei 'Milde, a slightly 
similar but nevertheless markedly distinct species; its label gives the locality etc. as: 
"Asplenium bourgaei, Turkey, Vil. Antalya (Isauria), Geyik Dag, 8000', crevices of 
shady limestone rocks, rare. P.M. Davis 1 451 3, 31 /8/1 947". It was with excitement 
that it was arranged for the specimen to be sent to Professor Reichstein for study as a 
possible candidate (at the time) for A daghestanicum, or at least as something 
different from other European and western Asian species. On seeing it he was able to 
suggest that it was close to but not the same as A daghestanicum and could well be a 
new species. However he suggested that there was also a possibility that it could be a 
distinctive but vicariant form of A creticum Lovis, Reichstein & Zaffran, a rare 
allotetraploid species from Crete derived from A viride and A aegaeum Lovis, 
Reichstein & Greuter, differing from it mainly in having the distinctive fused frond- 
apex. In A creticum the frond-apex is not normally fused but occasional small 
specimens can show some signs of slight fusion, though nothing like the extentthat it 
is fused in the Turkish specimen. The latter also has a narrower and less dissected 
frond than A creticum and the somewhat irregularly tripartite lower pinnae with the 
sori crowded near the base are again distinct. It was therefore reported as an 
Asplenium species near to (but distinct from) A creticum by Parris & Fraser-Jenkins 
(1980). 



164 FERN GAZETTE: VOLUME 1 3 PART 3 (1 987) 



In order to investigate the species further it was planned to visit the locality to 
search for more material, Professor Reichstein kindly agreeing to finance the 
excursion which was carried out in September 1 979 by the first-named author along 
with Miss S.J. de C. Coombs. 

PHYTOGEOGRAPHICAL NOTE 
Geyik Dag is part of the large area of high-level limestone shield comprising the Taurus 
Mountains (Toros Daglari) extending shortly inland all along the southern 
Mediterranean coast of Turkey and including some remote and very splendid 
mountains, often hard of access, such as Ak Dag (3073m) and Bey Daglari (3086m) in 
the western, or Lycian, Taurus, west of Antalya, another Ak Dag (c. 3100m), Bolkar 
Daglari (3240m). Aydos Dagi (3488m) and Toros Dagi (3585) in the central, or Isaurian, 
and eastern, or Cilician, Taurus east of Antalya and to the west and north of Adana. The 
shield also extends beyond the Taurus proper to Ala Daglari (391 Om)tothe north-east. 
These mountains can often be seen from a great distance as towering pale or white 
masses, not only because of their long- lasting snows, but also because of the very pale 
colour of their extensive and rugged limestone exposures. The Bey Daglari seentothe 
west from across the Gulf of Antalya (Antalya Korfezi), and the Bolkar Daglari seen 
from the north towering up above the dusty plain of Konya province (Vilhayet) are two 
of the best-known spectacular views of the Taurus. As one enters the mountains 
themselves north of Alanya, some 120km east of Antalya, the long high limestone 
ridge of Ak Dag (White Mountain) comes into view, just to the south-east of which lies 
the connected, sharp and rugged peak of Geyik Dag (2890m). Phytogeographically the 
Taurus Mountains lie within Mediterranean Turkey (see Parris and Fraser-Jenkins 
1 980), and form the border-line between that region and Inner Anatolia, but they are in 
many respects a special regionthemselvesduetotheir high altitude, and as far as both 
ferns and flowering plants are concerned they contain not only many endemicspecies 
but also a number of extensions of the ranges of European species, sometimes as a 
fragmented link between the flora of Greece (via the arc from the Peloponnese, 
through Crete and Rhodes, and thus to the western Taurus), and the flora of the 
limestone regions of the south-west Caucasus. An interesting example of this turned 
up during the trip concerned, with the surprise find of Dryopteris submontana (Fraser- 
Jenkins and Jermy) Fraser-Jenkins on the north side of Geyik Dag, new to Turkey (see 
Parris and Fraser-Jenkins 1980), and otherwise known from Europe, reaching its 
nearest point to Turkey at Mt. Parnassos in Greece. It is also known from the western 
Caucasus in Abkhazia, U.S.S.R., where the first-named author has been fortunate 
enough to see it in 1976; it was long known erroneously under the name D. villarii 
(Bell.) Woynar ex Schinz et Thell., as in Britain, before the taxonomy of the group was 
worked out (Fraser-Jenkins 1 977). The Turkish plants appeared to be slightly different 
from most of the European ones in their more adnate, less lobed pinnules with very 
long teeth at their apices (though similar plants also occur in Crna Gora (Montenegro), 
Jugoslavia, as, for example at the Cakor Pass, west of Pec), but they are clearly within 
the range of D. submontana and have the characteristic dense, sticky-glandularity of 
that species. A chromosome-count carried out by the second-named author on the 
plant from Geyik Dag (CRFJ 9809, BM) has confirmed that it is tetraploid (n = 82), 
which, along with its morphology, demonstrates its distinctness from the other two 
species in the erstwhile D. villarii cytological complex, D. villarii and D. pallida (Bory) 
C.Chr. ex Maire et Petitm., which are both diploid. Recently thefirst-named author has 
also seen a further, previously unidentified, specimen of D. submontana from the Ala 
Daglari (Nigde province), coll.: Find/ay 227 (E), and it could therefore also be expected 
elsewhere in the Taurus, including the Bey Daglari in the western Taurus. The 



ASPLENIUM TADEI 



165 



remoteness and lack of road access to many of the mountains in the Taurus range 
probably explains why it has not been found before and suggests the strong possibility 
of the existence there of other interesting and overlooked species of ferns. 




FIGURE 1. 

Holotype of Asplenium tadei 



COLLECTION OF THE NEW SPECIES 
On 9th September 1979 a first attempt was made to find the Asplenium, setting off by 
car well before dawn from a motel at Manavgat on the coast. The narrow and rough 
route to Geyik Dag turns off the main road north from Manavgat to Konya to head 
eastwards to the village of Gundogmus, in the foothills of the mountains. In roadside 
limestone crevices in the Pinus brutia Ten. forest there occurred at c. 750m alt. 
numerous populations of Dryopteris pallida subsp. libanotica (Ros.) Nardi, a diploid 
taxon very close to subsp. pallida, differing only in having distinctively marginal sori 
with a wide gap between the two rows of sori on each pinnule. Interestingly, in this 
area of Turkey there is a transition between subsp. pallida in the west and subsp. 
libanotica in the east (and Cyprus, the Lebanon etc.), and the populations here 
contained some clearly transitional forms (with good spores, not hybrids) as well as 
both subspecies. Beyond Gundogmus the forestry road turns northward into the main 
range and becomes even more narrow, rough and tortuous, being used mainly by 
small trucks sadly carrying away quantities of the natural forest as timber, and passing 
along spectacular cliff-edges above the forest, winding up into extremely rugged, 
higher- level ooen limestone crags, where it passes the shepherds' village of Guzel bag 



166 FERN GAZETTE: VOLUME 1 3 PART 3 (1 987) 



and comes out, some four hours (by Range Rover) after the start, at the base of the peak 
of Geyik Dag itself. On this first visit a direct and very hot and tiring ascent was made up 
the steep and dry west side of the mountain, over a ridge to the north-facing cirque, still 
containing snow-patches which last all through the summer, belowthesummit arete. 
At c. 2300m alt. plentiful Cystopteris alpina (Lam.) Desv. [= C. regia auct., non (L.) 
Desv.; lectotype in herb. Hort. Cliff. (BM!) is C. fragilis, (L.) Bernh., not Asplenium 
foreziense Le Grand as stated by Fuchs (1956 and 1 980)], with its very finely dissected 
fronds, was found, along with a hybrid, probably C. alpina x C. fragilis and also 
occasional Asplenium lepidum Presl. subsp. haussknechtii (Godot and Reuter) 
Brownsey, a species somewhat similar to A. ruta-muraria L.; but disappointingly there 
was no trace of the species being searched for. A second ascent was made on 1 2th 
September 1979 from the north side of the mountain after following a driveable 
shepherds' track around the base. Above seme shepherds' h-its at 2250m alt. a 
population of the correct Asplenium containing about 20 plants was discovered, with 
great delight, growing in crevices on some north-facing limestone cliffs. Unfortunately 
further investigation into the size and altitudinal extent of the population and into the 
presence of other species or hybrids suddenly became a great deal less interesting a 
proposition, and indeed was somewhat hastily abandoned due to the appearance of a 
gun-toting young man of somewhat wild appearance and ambiguous intentions, 
armed, as he demonstrated, with a fully loaded Czech pistol, and who, while waving 
his pistol in their direction, insisted on accompanying the collectors down the 
mountain in a state of slightly awkward tension to where the car was parked below. 
There, fortunately, traditional Muslim politeness was just ableto be maintained, with a 
few expressions of Turkish, and it was possible to turn the car and depart somewhat 
speedily around the side of the mountain and back on to the road-track. In all, five 
specimens of the Asplenium were collected and pressed (CRFJ 9812-9816) and an 
offset of CRFJ 9812 was transported living, partly wrapped in moss, to Basel to be 
grown by Professor Reichstein. It was immediately obvious in the field that this species 
did not closely match any other, including A. daghestanicum; it appeared in 
morphology most like A. viride, but with more deeply lobed lower pinnae and the 
distinctive fused frond-apex mentioned above. Although difficult to grow, including 
from fresh spores, the plant lived long enough in culture at Basel for a fixingto be made 
by Professor Reichstein and sent to the second-named author, who in August 1 981 , 
obtained a tetraploid chromosome count on it, with n = 72, at meiosis. In our opinion 
the collection clearly represents a new species, and we describe it, named in honour of 
Professor Tadeus Reichstein, thus:— 

DESCRIPTION 

Asplenium tadei Fraser-Jenkins et Schneller, sp. nov. 

Planta morphologia similaris ad A. viride sed pinnis profundiore lobatis et pinnatifidis- 
tripartitis earibus differt. Apex frondis insignis segmentum lanceolatum constantum ex 
aliquot paribus connatis pinnarum est et aliquot lobi angusti ad margines fert. Cytotypus 
tetraploideus sexualis. 

Holotypus: Turkey: "Asplenium tadei, sp. nov. Holotype. N. side of Geyik Dag, above 
shepherds' huts, N. of Gundogmus, N. of Alanya, Antalya Vilhayet, Turkey. N. facing 
shaded limestone cliffs, 2250m alt. C.R. Fraser-Jenkins (9815) and S.J. de C. Coombs, 
12/9/1979" (BM!)(Fig. 1). 

Isotypi: ditto (BM! Herb. T. Reichstein (5124 c), Basel!). Paratypi: ditto 981 2 (BAKU! CANV! 
BM! Herb. T. Reichstein (51 23 A), Basel!), 9813 (G! Herb. T. Reichstein (51 24 A), Basel!), 
9814 (K! P! PE! Herb. T. Reichstein (5124 B), Basel!) and 9816 (BM! Herb. T. Reichstein 
(5124 D), Basel!). Plant similar in morphology to A. viride but differing in its pinnae being 
more deeply lobed, with narrow extended lobes, and becoming pinnatifid-trilobate near the 
base of the lamina. Sori crowded near the base and centre of each pinna. The distinctive 
frond-apex is a lanceolate segment consisting of several fused pairs of pinnae and bears 



ASPLENIUM TADEI 



167 



several (c. 5-7) pairs of narrow lobes at the sides. Stipe often as long as the lamina, green, 
with a brown base extending up to y 3 of its length; bearing minute filiform, dark scales at its 
base which become smaller but are scattered up the whole length of the stipe. Cytotype 
tetraploid sexual (CRFJ 9812, det. J.J. Schneller; T. Reichstein, pers. comm. 18 August 
1981). 

ORIGIN AND RELATIONSHIPS 
As Asplenium tadei is a tetraploid species whose morphology does not fit any known 
diploid species it seems likely that it may be an allotetraploid (amphidiploid) species 
along with the great majority of other European and western Asian tetraploid species, 
and indeed the concept of a mixed morphology can probably explain most of the 
appearance of the species. From the narrowness of the frond, the undivided pinnae 
and mostly green stipe and rhachis it seems likely that A viride, with which it grows 
sympatrically on Geyik Da$, could very probably be one of its ancestral diploid species. 
The other ancestor is less clear, though it must be a species with more lobed and 
probably longer pinnae. As no other diploid species occur today in the European (sens, 
/at.) flora which have a fused frond apex similar to A tadei, it is impossible to do more at 
this stage than make a vague guess as.to the rest of its origin, and it should be borne in 
mind that there exist two groups of mainly Asian species, the A exiguum and A. 
daghestanicum aggregates which all have such a feature, so that A tadei may well 
belong to one of these groups as a markedly distinctive member, with a diploid 
ancester in one of the groups, possibly shared with A. daghestanicum. There is also a 
possibility that its second ancestor could be the south Turkish, Lebanese and Greek 
Aegean islands (Rhodes, Kastellorizo and Karpathos) endemic A bourgaei, another 
species present in the vicinity (see Greuteref a/. 1983), but at much lower altitudes (up 
to c. 900m) nearer the coast. A. bourgaei has longer, narrower pinnae bearing lobes 
which are widely joined at their bases; it also has atendencyforthe bases of the upper 
pinnae to fuse together near the frond apex, but not nearly as markedly or widely as in 
A. tadei. However, in an allopolyploid combination it is often difficult to predict which 
characteristics will be emphasised and which not. Greuter et a/. (1983) mentioned 
Davis' record of his specimen (which they did not see) under A. bourgaei, but stated 
correctly that because of its "completely aberrant" altitude it needed confirmation and 
might be due to some confusion. A. bourgaei has not so far been treated as in anyway 
close to A. exiguum, but we believe that both it and the rare and restricted south 
French endemic, A. jahandiezii (Litard.) Rouy are closer to the A. exiguum group 
(including the Sino-himalayan A. nesii Christ) than to any other European species, 
though on a larger, world-wide scale, such groups may not be so clearly applicable. 
Further study of/4, tadei including attempted hybridisation work will becarriedout 
by Professor Reichstein at Basel. 

ACKNOWLEDGEMENTS 
The authors wish to thank Professor Reichstein for his help in studying this species, 
and the first-named author is most grateful for his kind financial grant in aid of his 
expedition to collect it. They also wish to congratulate him on the achievement of his 
90th birthday. 



REFERENCES 
DAVIS, P.H. (ed.) 1965. Flora of Turkey and the East Aegean Islands, Vol. 1. Edinburgh. 
FRASER-JENKINS, C.R. 1977. Three species in the Dryopteris villarii aggregate (Pteridophyta, 

Aspidiaceae). Candollea 32: 305-319. 
FRASER-JENKINS, C.R., KHULLAR, S.P. & REICHSTEIN, T, in prep. (c. 1988). Pteridophyta, in 

Rechinger, K.H. (ed.), Flora Iranica. 



168 FERN GAZETTE: VOLUME 1 3 PART 3 (1987) 



FUCHS, H.P. 1956. Ph.D. thesis (Basel). 

FUCHS, H.P. 1 980. Beitrage zur Nomenklatur und Taxonomic der Schweizer Flora. Feddes Rep. 

90: 525-689. 
GREUTER, W., PLEGER, R., RAUS, T., ZIMMER, B. &GREUTER, J.J. \$Q3.Asplenium bourgaei, a 

new addition to the flora of Europe. Fern Gaz. 12(5): 271-274. 
PARRIS, B.S. & FRASER-JENKINS, C.R. 1980. A provisional checklist of Turkish Pteridophyta. 

Notes Roy. Bot. Gard. Edinburgh 38(2): 273-281. 






FERN GAZ. 13(3) 1987 169 



THE FERN HERBARIUM OF COL. F.J. HUTCHISON 

JOHN EDMONDSON 

Botany Department, Liverpool Museum, William Brown Street, 

Liverpool, L3 8EN 

ABSTRACT 

The history and composition of F.J. Hutchison's fern herbarium is given, based on a 
manuscript catalogue prepared by H. Stansfield, and extracts are quoted from 
relevant correspondence. The herbarium contains material from 24 collectors, 
including Edwin Atkinson, Lady Anne Barkly, Francis Brent, John Buchanan! 
Charles B. Clarke, John Day, Wilhelm Hillebrand, George F. Hose (Bishop of 
Singapore & Sarawak), F.J. Hutchison and his probable relative H. McLeod 
Hutchison, Phoebe Moss, Charles Parish, Thomas Powell, Richard Spruce, William 
Stout and George Wall. Hutchison's own collections were largely made between 
1 870 and 1 872 in Ceylon (Sri Lanka); they came to Liverpool Museum via the Royal 
Albert Memorial Museum, Exeter. 

HISTORY OF THE COLLECTION 
One of the largest collections of tropical ferns in the herbarium of Liverpool Museum 
(LIV) was presented in 1947 by the Royal Albert Memorial Museum, Exeter. It had 
been acquired from the widow of Col. Frederick J. Hutchison, shortly after his death on 
December 2, 1 981 at Stoke, Devonport, Devon. The collection amountstosome 1,800 
mounted sheets, and is noteworthy because it is composed not only of material 
collected by Hutchison himself in Ceylon, but also of exchange herbarium material 
sent by some well-known pteridologists of the mid-Victorian period. 

Three categories of material can be recognised. Firstly, there are specimens 
which Hutchison himself collected between 1870 and 1872 in Ceylon (and a few from 
Dartmoor and Bovisands (1875), Perthshire (1876) and Ascot (1879). The total number 
comes to 31 sheets. The principal collection localities in Ceylon are given in Table 1 . 

Secondly, specimens acquired by Hutchison from his pteridological contacts 
amount to some 1 553 sheets. A list of these collectors is given below. Both categories 
consist of material mounted on "fern-size" sheets (21 x 13 inches), are fully 
catalogued and numbered in taxonomic sequence from 1 to 1,863. 

Lastly, there is a large and bulky series of unmounted material, much of which is 
only skimpily labelled. Some may duplicate Hutchison's collections. The number of 
unmounted and uncatalogued specimens was estimated at 9,000; this figure should 
be treated with caution, as some of the material consists of sequences of dismembered 
frond segments stored in separate paper folders which could be reassembled into a 
single frond. 

The catalogue of the mounted material in the Hutchison fern herbarium was 
prepared by the Museum's former Keeper of Botany, Mr H. Stansfield, and was 
completed in 1 958. No attempt was made to revise the nomenclature, which retains 
the taxonomic treatment adopted by Hutchison. This adheres closely to Synopsis 
Filicum (Hooker & Baker, 1 974). Many of the statistical data in this article have been 
extracted from Stansfield's catalogue. 

TABLE 1. 

Gazetteer of collection localities in Ceylon (Sri Lanka) visited by F.J. Hutchison 

(Spelling as on labels) 



Aggelwatte 


10.1871 


Galagana 


4.1872 


Bullatotte Pass 


7.1871 


Galle 


7.1871 


Colombo 


8-9.1870, 7.1871, 9.1871 


Gonaganna 


2.1872 


Condagalle Pass 


1.1871 


Gongolla 


4.1872 


Dimboola Pass 


3.1871 


Hakgalla 


1-2.1871, 5.1871 



170 



FERN GAZETTE: VOLUME 13 PART 3 (1987) 



Hantam 


1.1871 


Hapootella 


5.1871 


Helbodde 


8.1870 


Hewissa 


10.1871 


Hoolakande Pass 


8.1871 


Horton Plains 


4.1872 


Kambodde 


8.1870, 1 


Kandy 


10.1870 


Kegalli 


6.1872 


Kornegalle 


2.1870 


Maturalta 


4.1871 


Morowa Korle 


7.1871 



2.1871 



Newera Enya 


1-2.1871 


Oodoowelle 


10.1870, 1.1871 


Pittawelle 


8.1871 


Poorie 


5.1872 


Puselawe 


8.1870, 1.1871 


Rambodde Pass 


1.1871 


Rambodde 


2.1871 


Rottowa forest 


7.1871 


Telgamma 


10.1871 


Vicarton 


8.1871 


West Matale 


8.1871 



HUTCHISON'S FELLOW ENTHUSIASTS 
Although Hutchison's own collecting activities in Ceylon were carried out privately 
while he was stationed there (with the rank of Major), it is clear from correspondence 
held at Liverpool Museum that he collaborated closely withthe principal pteridologists 
of the period in Ceylon. One of these was Thomas W. Naylor Beckett (1839-1906), a 
coffee planter, whose herbarium and correspondence is kept at Liverpool Museum. 
Naylor Beckett was born in Liverpool and spent much of his life in Ceylon before 
moving to New Zealand in 1883. 

" A letter addressed to Naylor Beckett from George Wall, a fern enthusiast resident 
in Ceylon, describes a recently completed fern-hunting expedition. Dated "Colombo, 
July 23, 1871", it commences: 

"My dear Sir, 

I returned here last night from a trip through Moosoowakka Suffragum & Rahnapoora with 
Major Hutchison. I went on business but took my fern papers and worked on the way. Your kind 
letter and most valuable catalogue awaited my return and I lose notime inthankingyouforthem. 

We had awful weather, but except on one day we carried out our programme and worked 
through the dreadful downpour, regardless of leeches and all the multiform discomforts of 
incipient rain. — I think we have made a very good bag, all things considered, and we have 
confirmed an opinion which we have both gradually been brought to that even the rare ferns are 
much more widely spread than generally supposed. Your list is in itself a strong confirmation of 
that opinion too ..." 

The letter continues with a list of the chief new finds. It was written on a date 
midway through Hutchison's period of fieldwork; see Table 1. 

Included in the correspondence on file in Liverpool is a letter to Beckett from F.J. 

Hutchison dated "Columbo, 30 Dec. '71". It reads, in part: 

"... Mr Wall wrote to you a full account of our doings at Matale, and I hope expressed, as I asked 
him to, my acknowledgments to you, with his own, for the excellent instructions you have given 
us, thanks to which the expedition was the most successful we had ever made ... We had, as you 
know, planned two other expeditions, one to your part of the country and another to the Singh 
Raja Forest, both of which Mr Wall's engagements forced him to give up, much to our 
disappointment. The latter I have since partially accomplished, partially only, because Ifoundthe 
distances were not at all what I had been led to expect. In fact to quote Mr Thwaites, to whom I 
sent an account of my doings, I got "the cream of the Singh Raja Forest ferns." 

Mr Wall has written to methricesince he reached England. He has had some interviews with 
Baker at Kew[1], and sent me a copy of notes made thereon some of the Ceylon ferns which I will 
enclose as I have no doubt you will be interested in them. You will seethat Baker pronounces the 
Acrostichum, which we found near Morawaka, and of which Mr Wall sent you a specimen, to be a 
distinct species [2] and not merely a form of A. (Gymnopteris) variabile as we supposed ... 

There will not I fancy be many gaps in your Ceylon collection but it is possible that I may be 
able to fill some of them for you, and I need scarcely say that it will give me the greatest pleasure to 
do so ..." 

Notes: [1] John Gilbert Baker (1834-1920) was then an Assistant Curator of the Kew 
herbarium; later (1890-1899) he held the position of Keeper. 

[2] Possibly a reference to Acrostichum wallii Baker, ascribed in the Appendix to Hooker & 
Baker (1874), p. 523. 






COL HUTCHISON'S FERN HERBARIUM 171 



There is also at Liverpool a letter to Beckett from Wall dated 'Colombo, 1 9 July 
1872" which refers to "Major H.". The text of the letter is as follows: 
"My dear Sir, 

I acknowledged your kind letter a day or two ago and I now send you a list of the ferns I have 
yet to find. It is cleared of several by my last trip and does not contain a large proportion of very rare 
ones. Of these I have been fortunate in finding a good many. Of Diacalpe Aspidioides for instance 
Major H. and I discovered the headquarters between Mathooratee[1]and Ordupusselmonon the 
Kooroonda Oga where we found it in profusion and of great size. One frond I gave MrThwaites[2] 
was 2 feet long without the stipes. — Still I have a deal to do to complete my collection of known 
ferns, especially as some of those I found at first I spoiled not knowing how to manage them, i 
have not more than 150 or 160 species mounted and fit to mount. 

I have also suffered loss by insects, and it is a serious labour to poison all even with my rolling 
apparatus, which is infinitely more expeditious than the feather or brush. I hear there is a better 
method of poisoning by means of the fumes of Carbolic Acid. — Do you know this method? 

I fear I shall not be able to visit Dick[3] or you again so soon as I intended: but I must go to East 
Matali and to Singhe Rajah forest before I go home if possible - 

The Nephrodieae trouble me so much and I do not take much notice of them, as I expect to 
tackle them separately when I have cleared off all the rest. Latterly I have collected considerable 
quantities of all the ferns I have found, sol hope to be able to requite your kindness by contributing 
to your collection. I send herein a frond of an Acrostichum which I suppose I am bound to call Ac. 
Variabile [41 but my series of this plant was already most remarkable before I added this to the 
strange variety of forms this fern assumes. Do you know this form?- 

More anon 

Yrs very Sincerely, 
George Wall." 
Notes: [1] Hutchison has a specimen of Diacalpe aspidioides (cat. no. 103) from "Maturatte, April 
71". This species is now sometimes known as Peranema aspidioides (Blume) Mett. (e.g. Sledge, 
1982). 

[2] G.H.W. Thwaites (1 81 2-1 882) was, at thetime, Director of the Botanic Gardens at Peradeniya, 
Ceylon. According to Desmond (1977), G. Wall was a friend of his. 

[3] "Dick" is possibly a reference to Richard Henry Beddome (1839-191 1), author of "Ferns of 
British India" (1865-70). 
[4] Now known as Leptochilus decurrens Blume. 

The Hutchison herbarium contains 42 specimens contributed by George Wall in 
addition to those which were collected during joint excursions. It is significant, 
however, that Hutchison did not receive material from either Thwaites or Naylor 
Beckett. 

The list of collectors represented in the Hutchison herbarium includes only one 
other botanist resident in Ceylon: William Ferguson (1820-1887), a civil servant in 
Ceylon who published various works including "Ceylon Ferns" (1872) and "Ceylon 
Ferns and their Allies" (1880). There are letters from Ferguson to Naylor Beckett in the 
files at Liverpool, one of which reveals that Beckett assisted Ferguson in the editing of 
his "Ceylon Ferns" prior to publication. 

The picture one builds up is of a group of enthusiasts co-operating in the mutual 
enrichment of their collections, some of whom — Hutchison included — were able, 
through fieldwork, to contribute rarities at the request of other collectors and who in 
turn acquired valuable exsiccata from other parts of the world. 

INDEX OF COLLECTORS 
It is unclear how Hutchison obtained material from collectors of non-Ceylon material 
and if information comes to light the Liverpool Museum would be grateful to receive it. 
The following collectors are the principal outside contributors to the Hutchison 
herbarium (numbers of specimens are given in square brackets): 
Atkinson, Edwin (1840-1890). Entomologist, Indian Civil Service, 1862-1890. 
Sikkim, Khasya Hills, East Indies. [50] 

Barkly, Lady Anne Maria (1838-1922). Second wife of Sir H. Barkly (1815-1898). 
Reunion, Mauritius, Namaqualand (Namibia, S.W. Africa). [44] 

Boyd, Miss. Dates and occupation unknown. Sandwich Is. (Hawaii). [15] 

Brent, Francis (1816-1903). H.M. Customs officer; the specimens are from Brazil, 
though it is unlikely that Brent collected them himself. There are no references 
to Brent, nor to Randall (q.v.) as collectors in Martius & Eichler, Flora Brasiliensis 



172 FERN GAZETTE: VOLUME 13 PART 3 (1987) 



vol. 1, 1906: chapter entitled "Vitae Itineraque Collectorum Botanicorum" 

See also Greenwood (1 972). [16] 

Buchanan, Revd. John. Minister, Durban, South Africa, 1861-1874. 

Orange River Colony, South Africa. [21 1] 

Clarke, Charles Baron (1832-1906). Superintendent of Calcutta Botanic Gardens, 

1869-71; worked at Kew after retirement. Co-author (with Henderson) of 

"Ferns of N. India" (1880). Dhurmsala, India, 1874. [1] 

Corrie, Dr. Dates and occupation unknown. Fiji. [few] 

Craig, ?William (fl. 1880-1890), an Auckland dealer in fern exsiccata. 

Provenance given as "New Caledonia". Although only a few specimens are 

attributed to "Craig" in Stansfield's catalogue, there is a significant number of 

specimens from New Caledonia which lack details of the collector. By checking 

the names of such ferns in Flore de la Nouvelle Cale'donie. no. 3: Pte'ridophytes 

(Brownlie, 1969) it is clear that they include a number of endemic species 

collected by Vieillard. It is likely, though difficult to prove, that Craig supplied 

Hutchison with a collection whose labels had been lost or discarded. 

Day, John (1824-1888). Amateur fern and orchid enthusiast and traveller. 

Australia, Ceylon, Fiji, Indonesia, Japan, New Zealand, West Indies 

(Trinidad, Jamaica). Most are from Japan. [252] 

Ferguson, William (1820-1887). Surveyor, Ceylon Civil Service. Author of 

"Ceylon Ferns". Ceylon, St. Helena. • [3] 

Gamble, James Sykes (1847-1925). Employed in the Indian Forest Service 

from 1871, and Director of the Forestry School at Dehra Dun from 1890-99. 

Sikkim, 1872. [2] 

Grant, James Augustus (1827-1892). Army Officer. Hong Kong. [1] 

Gray, Samuel O. (1828-1902). Author of "British Seaweeds" (1867). 

West Indies (St. Vincent). [2] 

Hillebrand, Wilhelm (1821-1886). German doctor and botanist, resident in Hawaii 

1851-1871. Author of "Flora of the Hawaiian Islands" (1888). 

Java, Samoa, Sandwich Islands (Hawaii). [114] 

Hose, Revd. George Frederick (1838-1922). Bishop of Singapore and Sarawak, 

1881-1908. Malaya, Sarawak. [83] 

Hutchison, H. McLeod (1840-1925). Army Officer. Probably a relative of 

F.J. Hutchison, but exact relationship unknown. According to Hart's Army List 

for 1867, he was Lieut, in the 14th (Buckinghamshire) Regiment of Foot, whose 

2nd Battalion returned from Jamaica in 1 864. Jamaica (see also Prior). [119] 

Moss, Miss Phoebe (dates unknown). An amateur naturalist who, in 1885, 

introduced the Common Myna (Acridotheres tristis) and the Grass Frog (Rana grayi grayi) 

to St. Helena. [Information from Dr Q. Cronk in litt. 24. 1 1 . 1 982]. 

St. Helena, South Atlantic Ocean. [1 1] 

Munroe, W. (1818-1880). Army Officer and agrostologist; General, 1878-1880). 

Canada (Quebec), 1859. [2] 

Parish, Revd. Charles S.P. (1822-1897). Army Chaplain. Burma (Moulmein). [42] 

Powell, Revd. Thomas (1809-1887). Missionary in Samoa, 1860-85. Samoa. [29] 

Prior, Richard C.A. (1809-1902). Curator of Fielding Herbarium, Oxford. Jamaica. 

Many Jamaican specimens lack details of the collector; they may have been collected 

by either H. McL. Hutchison or Prior. 

Randall, Mr. Dates unknown. Brazil. See note under Brent. [20] 

Spruce, Richard (1817-1893). One of the most notable botanical explorers 

of South America. Mainly Peru. [11] 

Stout, William (d. 1882). U.S.A. According to Joseph Ewan, this is one of the largest 

collections of Stout's material. [152] 

Wall, George (c. 1821-1894). Head of the firm of George Wall & Co., a businessman 

based in Colombo and keen amateur pteridologist. Ceylon. [42] 

ACKNOWLEDGEMENT 
I am indebted to Stephen Harrison for his unpublished notes on the Hutchison 
herbarium, which saved me much time in the preparation of this account. 

REFERENCES 
BEDDOME, R.H. 1865-1870. The ferns of British India. Madras. 
BROWNLIE, G. 1969. Flore de la Nouvelle Cale'donie, no. 3. Pte'ridophytes. Paris. 
DESMOND, R. 1977. Dictionary of British and Irish Botanists and Horticulturalists. London. 
GREENWOOD, E.F. 1972. The Beans of Scarborough. J. Soc. Bib/. Nat. Hist. 6(3): 152-161. 
HOOKER, W.J. & BAKER, J.G. 1874. Synopsis Filicum, 2nd edition. London. 
SLEDGE, W.A. 1 982. An annotated check-list of the Pteridophyta of Ceylon. Bot. J. Linn. Soc. 84: 
1-30. 



FERN GAZ. 13(3) 1987 173 



THE STATUS OF OPHIOGLOSSUM AZORICUM 

(OPHIOGLOSSACEAE : PTERIDOPHYTA) 

IN THE BRITISH ISLES 

A.M. PAUL 

Botany Department, British Museum (Natural History), Cromwell Road, 

London, SW7 5BD, England 

ABSTRACT 

The status of Ophioglossum azoricum in the British Isles is examined by comparing 
the gross morphology, epidermal features, spores and cytology of the three British 
species of Ophioglossum, O. azoricum, 0. lusitanicum and O. vulgatum, together 
with their ecology and distribution. The possibility that O. azoricum is just an 
ecotype of O. vulgatum is considered. It is concluded that these two taxa cannot be 
satisfactorily separated morphologically and that 0. azoricum appears to be merely 
at one end of the spectrum of variation shown by O. vulgatum. 0. azoricum is 
maintained as a separate species for the present because certain populations are so 
distinct in habit, but it is now evident that cytological investigation combined with 
extensive field studies is necessary to elucidate the status of this taxon. 

INTRODUCTION 
The taxon now known as Ophioglossum azoricum C. Presl was first found and 
recognised in Britain as distinct by J. Boswell Syme in the Orkneys in the mid- 1 9th 
century (Boswell Syme 1871). Due to its similarity to 0. vulgatum L, it has been 
referredto as a variety or subspecies of that species inthe European literaturefor well 
over 100 years. However, in 1964, Rothmaler accepted 0. azoricum as a species. He 
described it as morphologically distinct from O. vulgatum, and intermediate in many 
respects between that species and 0. lusitanicum L. 

The nomenclature of the taxon has been rather confused (see synonymy below). 
This can be principally attributed to the existence in Africaand parts of western Asiaof 
a further taxon, O. polyphyllum A. Braun, which was erroneously regarded by some 
botanists as a subspecies or variety of 0. vulgatum, and to which some European 
specimens were referred. Pichi-Sermolli (1954) showed that 0. polyphyllum was 
distinct from the European plants, which he referred to O. vulgatum var. ambiguum 
Coss. &Germ. A comparison of these two taxa and their distribution in Macaronesia is 
given by Lobin (1986). 

There are three species of Ophioglossum in the British Isles, of which one, 0. 
lusitanicum, may easily be distinguished morphologically from the other two. Its 
winter growing season also means its identity is unlikely to be mistaken. 0. azoricum 
and 0. vulgatum, however, are less easy to separate, as most of their characters 
overlap markedly in range. This comparative study ofthethreetaxa was undertaken to 
find characters to distinguish the two summer-growing species, and also to see if 0. 
azoricum is truly intermediate between the other two and sufficiently distinct to 
warrant specific recognition. 

Synonymy 

Ophioglossum azoricum C. B. Presl. 1845. Suppl. Tent. Pterid.: 49. 

0. vulgatum subsp. polyphyllum E.F. Warburg. 1 952. in Clapham, A.R., T.G. Tutin &ER 
Warburg, Fl. Brit. Is.: 54. pro parte excl. syn. A. Br. 

O. vulgatum subsp. ambiguum (Coss. & Germ.) E.F. Warburg. 1957. Watsonia. 4: 41. 
0. vulgatum var. vel subsp. polyphyllum auct.; non 0. polyphyllum A. Braun in Seubert, 
1844. Fl. Azor.: 17. (vide Pichi-Sermolli 1954). 



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FERN GAZETTE: VOLUME 13 PART 3 (1987) 



CYTOLOGY 
Love and Kapoor (1966, 1967) suggest that 0. azoricum is of hybrid origin, an 
allopolyploid derivative of a chance hybrid between O. vulgatum and 0. /us/tan/cum. 
The taxon may be intermediate morphologically, but the cytological basis for their 
hypothesis is founded on insubstantial evidence. 

Few chromosome counts have been reported for European specimens of 
Ophioglossum. These are summarised in Table 1. As yet, counts for British plants have 
been published from only one sample each of O. /us/tan/cum andO. vulgatum {Manton 
1950) and none for 0. azoricum. European counts suggest the chromosome number of 
0. vulgatum is n = 240-260, and of 0. lusitanicum is n = 1 25-1 30. However, Love and 
Kapoor (1 966, 1 967) give the chromosome number of O. lusitanicum as 2n = 240, but 
with no reference source. Furthermore, their count for O. azoricum (2n = 720) is based 
only on Icelandic material. They consider their earlier count (given under the name O. 
vulgatum ssp. ambiguum var. islandicum Love & Love, in Love & Love 1961) of 2n = 
480 to be an inexact estimate. This is presumably the basis of '2n = c.480' cited for O. 
azoricum in Flora Europaea; Rothmaler's decision to recognise the taxon here as a 
good species was not based on cytological evidence. 

Further cytological studies of this genus in Britain and Europe as a whole 
(including the Azores) are obviously required, particularly to help establish the status 
of 0. azoricum. It is interesting to note that Japanese material of O. vulgatum has also 
given a chromosome count of n = 240 (Kurita & Nishida 1965), but that Indian 
specimens of O. lusitanicum and 0. vulgatum have revealed chromosome numbers of 
a range approximately double those of European and Japanese material (Ninan 1 956, 
1958, Verma 1956). 





TABLE 1. European Ophioglossum chromosome 


counts 




Species 


Origin 


Author & Date 


n 


2n 


0. lusitanicum 


Guernsey 


Manton 1950 


125-130 
c. 128 




0. vulgatum 


England 


Manton 1950 


250-260 

c. 256 




0. vulgatum 


Sweden 


Ehrenberg 1945 




c. 344* 


0. vulgatum 


Netherlands 


Verma 1958 


240, 247- 
251 (c. 248) 




0. vulgatum 


Finland 


Sorsa 1962 


c. 250 




0. vulgatum 


Spain 


Love & Kjellqvist 
1972 




c. 480 


0. vulgatum 


Sweden 


Love & Love in 
Love 1976 




480 


0. azoricum** 


Iceland 


Love & Love 1961 




480 


0. azoricum 


Iceland 


Love & Kapoor 










1966, 1967 


360 


720 






Love & Love 










in Love 1976 




720 



based on sectioned cells rather than a 'squash' 
**as O. vulgatum subsp. ambiguum var. islandicum 



MORPHOLOGY 
General morphology 

As previously mentioned, 0. lusitanicum is morphologically distinct from the other two 
British species, but O. azoricum and 0. vulgatum overlap considerably in their range 
for most characters. They are usually separated by the number of fronds per plant, 



THE STATUS OF OPHIOGLOSSUM AZORICUM IN THE BRITISH ISLES 



175 




FIGURE 1. a, Ophioglossum vulgatum, Morfa Dyffryn, Merioneth. x1. (photo. A. Cleave), b, 0. 
lusitanicum, St. Agnes, Scilly Isles. x2. (photo. K.H. Hyatt), c, 0. azoricum. Ravenglass, 
Cumberland. x2. (photo. A.C. Jermy). d, 0. azoricum, S. Uist, Outer Hebrides. x2. (photo. A. Cleave). 



176 



FERN GAZETTE: VOLUME 13 PART 3 (1987) 



frond size and shape, and the number of sporangia per fertile spike. For example, two 
of the criteria used to recognise O. azoricum for the Atlas of Ferns of the British Isles 
(ed. A.C. Jermy et al. 1978) were 14 or less pairs of sporangia and a sterile blade less 
than 3.5cm long. 

0. vu/gatum is said to produce usually only one frond, occasionally two per plant, 
whilst O. azoricum and 0. lusitanicum regularly bear 2-3 fronds. However, 
populations observed in the field or preserved as herbarium specimens indicatethat 0. 
vu/gatum produces more than one frond more commonly than is generally supposed, 
andthe other species have usually two fronds, sometimes one, andoccasionallythree. 

O. lusitanicum is not a variable species. Sterile blades are narrowly lanceolate, 8- 
23(-38)mm long, 1.5-4(-6)mm wide. They are widest at about the centre of the frond, 
narrowing gradually to the base and apex, the latter being slightly rounded. The sterile 
blades of 0. vu/gatum tend to be broadly ovate-lanceolate to ovate, usually widest 
below the middle. They are extremely variable in size and shape even within a single 
population, 17-150 x 8-58mm, with the apex ranging from acute to very rounded or 
apiculate. The base may be attenuate to cordate, often clasping the stem. Some 
specimens of 0. vu/gatum, on the other hand, are relatively small, with lanceolate 
sterile blades, and are very similar to those of O. azoricum. The latter have blades 9-22 
(-40) x 4- 1 0(- 1 4)mm which are broadly lanceolate to ovate, usually with cuneate bases 
and acute to obtuse apices, the lamina being broadest at or just below the centre. 




FIGURE 2. Herbarium specimens of Ophioglossum. a, O. vu/gatum, R. Vowell Sherring, 1912, 
Enfield, Middlesex. x0.5. b, 0. lusitanicum, M. Dawber, 7 January 1886, Guernsey, Channel 
Islands. x1. c, 0. vu/gatum, R.J. Pankhurst & A.O. Chater, 12 August 1983, Berneray, Outer 
Hebrides. x1. d, 0. azoricum, A. Cleave, June 1983, S. Uist, Outer Hebrides. x1. 



THE STATUS OF OPHIOGLOSSUM AZORICUM IN THE BRITISH ISLES 177 



The angle at which the sterile blades grow is sometimes distinctive for different 
Ophioglossum species. Those of 0. lusitanicum are often reflexed, and so held close to 
the ground (Fig. 1 b), though occasionally they are more upright. Fronds of 0. vu/gatum, 
by contrast, tendto be quite erect (Fig. 1a). O. azoricum tends to resemble the former in 
its reflexed habit (Fig. 1c, d), a feature not mentioned in thetype description, but shown 
by some individuals of the isotype (BM). This is, of course, a character which is often 
lost on herbarium specimens, however carefully they are pressed (Fig. 2), but it seems, 
generally speaking, to be one of the best field characters. 

The other principal character commonly used is the number of pairs of sporangia 
on the fertile spike. O. lusitanicum has 3-8(-1 3), O. vu/gatum 1 1-44, and O. azoricum 
4- 1 3(- 1 7), the range for the latter overlapping that of the other two species. Size of 
fronds and sporangial number do not seem to be directly correlated; some specimens 
of 0. vu/gatum with tiny leaves (less than 2cm long) may have as many as 20 
sporangial pairs, others far fewer, and likewise some large individuals have relatively 
few sporangia. 

Other features of plants, such as the length of the fertile spike and the common 
stalk, and the relative proportions of fertile blades were studied for a large number of 
specimens, but did not prove to be useful in identification. They are very varjable, 
differing with the maturity of the plant, depth of the rhizome below the soil surface, 
habitat and climate. The shape of the rhizome is also not constant. 0. lusitanicum does 
tend to have an elongated, rather cylindrical rhizome, but in the other species small 
specimens have relatively small, globose rhizomes, whereas in larger individualsthey 
are considerably bigger and much more elongated. Hand-cut sections of roots of 
varying sizes stained with safranin showed the stele of all three taxa to be monarch. 

Venation and epidermal characters 

Venation was one of the key characters on which Prantl (1 884) based his classification 
of Ophioglossum. Clausen (1 938), however, realised that venation was not a reliable 
character, due to its variability within a single population. In 1 962, Mahabale based a 
key to Indian Ophioglossum species on characters of venation. Panigrahi and Dixit 
(1969) likewise considered venation important in identification, and Bhambie and 
Madan (1982) also reported venation patterns to be taxonomically significant at 
species level, in terms of areole number per unit area of frond. However, Wagner et at. 
(1 981 ) and Wagner et al. (1 984) showed that a single species could produce a range of 
fronds, from small ones with simple venation to large forms with more complex 
venation. 

In this study, fronds taken from herbarium sheets were cleared in 10% bleach 
('Domestos') and stained with an aqueous solution of methylene blue. They were 
thereafter stored in 50% alcohol. Venation patterns were drawn using a microscope 
drawing attachment, or by tracing photographs. 0. lusitanicum was the most distinct 
species, having only a primary network of veins forming rather long, narrow areoles 
parallel with the mid-vein. Very few included free veinlets are present (Fig. 3a, b). The 
areoles of the primary vein reticulum of 0. azoricum are less protracted, and the more 
central, larger areoles contain a finer secondary mesh. Included free veinlets are 
common but not numerous (Fig. 3c, d). The venation pattern of 0. vu/gatum is much 
more variable, probably owing to its range of frond size. In large fronds, the majority of 
primary areoles contain finer veins forming a secondary network, and also many free 
included veinlets (Fig. 3f). In smaller blades the secondary reticulum is less well- 
developed and has few free included veinlets (Fig. 3e), thus not being very significantly 
different to that of O. azoricum. Since it is usually small-fronded 0. vulgatum 
specimens which tend to be confused with specimens of 0. azoricum, it appears that 



178 



FERN GAZETTE: VOLUME 13 PART 3 (1987) 




FIGURE 3. Venation of Ophioglossum. a, b, d, central portion across whole width of frond, c, e, f , 
central portion of frond from mid-vein to margin, a, O. lusitanicum, J.E. Gardner, Guernsey, 
Channel Islands, b, O. lusitanicum M. Dawber,21 January 1886, Guernsey, Channel Islands, c, 
0. azoricum, A. Cleave, June 1 982, Lundy, Devon, d, 0. azoricum, J. Boswell Syme, August 1 873, 
Swanbister, Orkney, e, O. vulgatum, R.J. Pankhurst & A.O. Chater, 12 August 1983, Berneray, 
Outer Hebrides, f, O. vulgatum, LH.J. Williams 1053, 12 June 1935, Kemsing, Kent. 



THE STATUS OF OPHIOGLOSSUM AZORICUM IN THE BRITISH ISLES 179 



venation characters are of rather limited usefulness. If extensive studies of fronds of 
different sizes and developmental stages could be undertaken similar to those of 
Wagner et al. (1981), differences between the venation patterns of O. azoricum and 
small individuals of 0. vu/gatum might become evident. However, it is equally possible 
that 0. azoricum would prove merely to be at one end of the spectrum of variation 
shown by O. vu/gatum. 

Epidermal characters, particularly cell-shape and stomatal orientation, have aiso 
been considered useful in distinguishing species of Ophioglossum (Prantl 1884, 
Mahabale 1962, Maroti 1965, Van Cotthem 1970, Pant & Khare 1969). For this 
investigation whole fronds, cleared and stained as above, were mounted and 
photographed using bright field microscopy. In one case the epidermis was peeled off 
and mounted in glycerine jelly. In each sample the area between the mid-vein and 
margin in the centre of the frond was photographed for comparative study and then 
drawn from the negative with the aid of a photographic enlarger. 

Once again it is O. /us/tan/cum which is markedly different from the other species. 
The epidermal cells on the lower surface are narrowly rectangular and parallel with 
the long axis of the frond (Fig. 4b); on the upper surface the cells aresquarer and more 
rounded (Fig. 4a). The lower epidermal cells of large fronds of 0. vu/gatum have very 
undulating margins, tending to be fairly long and narrow and parallel with the long axis 
of the frond (Fig. 4h); those of the upper surface have similarly undulating edges, but 
they are much more irregularly shaped (Fig. 4g). In small-fronded individuals, the 
epidermal cells of both surfaces are much smaller, with very much less sinuous 
margins (Fig. 4e, f). 0. azoricum specimens studied greatly resemble small individuals 
of O. vu/gatum in the shape of epidermal cells though the margins are rarely sinuous 
(Fig. 4c, d). 

In all three species the stomata are sunken, and distributed more or less equally 
on both surfaces of the fronds. On the lower surfaces the stomata are all parallel with 
the main vein (Fig. 4b, d, f, h). In 0. lusitanicum, the stomata on the upper surface are 
also mostly parallel with the long axis of the frond (Fig. 4a), but in the other two species 
they are randomly orientated (Fig. 4c, e, g), even in the centre of the frond. 

Thus it seems that the epidermis does not provide good characters for separating 
0. azoricum and small specimens of 0. vu/gatum. 

Spores 

As early as 1857 spores were considered diagnostic for distinguishing European 
Ophioglossum species (Durieu de Maisonneuve 1857). Prantl (1884) placed 
considerable emphasis on spore size and ornamentation in his classification of the 
genus and these characters were also found to be useful in distinguishing Japanese 
species (Nakamura & Shibasaki 1959, Kurita 1981) and Indian species of 
Ophioglossum (Mahabale 1962, Pant & Khare 1971). However, in the most recent 
monograph of the genus, Clausen (1 938) found spore characters to be of only limited 
value in distinguishing species, although this may in part reflect his broad species 
concept. Wieffering, in his preliminary revision of Indo-Pacific species of 
Ophioglossum (1964) implied that spores were characteristic only of sub-genera 
rather than species. 

All spores used for this study were removed from herbarium specimens and were 
not chemically treated prior to observation. Spores were glued to stubs with 'Araldite', 
coated with gold palladium and examined with an Hitachi S800 SEM. Permanent 
preparations are kept at BM. 



180 



FERN GAZETTE: VOLUME 1 3 PART 3 (1987) 




FIGURE 4. Epidermal features of Ophioglossum. Portion illustrated is from centre of sterile blade 
half-way between mid-vein and margin; vertical axis of frond is horizontal across page. a,b, O. 
lusitanicum, G. Wolsey, March 1 869, Guernsey, Channel Islands; a, upper epidermis and b, lower 
epidermis. c,d, 0. azoricum, A. Cleave, June 1 982, Lundy, Devon; c, upper epidermis and.d, lower 
epidermis. e,f, 0. vu/gatum, R.J. Pankhurst & A.O. Chater, 12 August 1983, Berneray Outer 
Hebrides; e, upper epidermis and f, lower epidermis. g,h, O. vu/gatum, F.J. Hanbury, 12 June 
1875, Trotter's Cliff, Kent; g, upper epidermis and h, lower epidermis. 



THE STATUS OF OPHIOGLOSSUM AZORICUM IN THE BRITISH ISLES 181 



The equatorial diameter of spores, taken from at least five herbarium specimens of 
each species, was measured in air. The overall size range for 30 spores from each 
sample was as follows (range of sample means in bold):— 0. vulgatum 28-30-38- 
43um, 0. lusitanicum 28-29-34-40um, 0. azoricum 35-38-42-45um. Page (1982) 
also found that spores of O. lusitanicum tend to be smaller, and those of O. azoricum 
larger, than O. vulgatum spores, but Ferrarini et al. (1986) report spores of 0. 
lusitanicum in Italy to be intermediate in size between those of 0. vulgatum and 0. 
azoricum. However, the ranges overlap considerably, and since a single sample may 
have a range of up to 1 0um, it is important to use the mean size of a large sample. It 
should also be noted that the medium in which spores are measured has a bearing on 
size, and it is advisable to use controls of known identity. 

Spore size is thus of limited usefulness, and the same seems to be true of the 
ornamentation of spores, as revealed by light microscopy (LM) and scanning electron 
microscopy (SEM). Spores of European 0. vulgatum have been variously reported as 
being tuberculate (Durieu de Maisonneuve 1 857, Rothmaler 1 964), having small, but 
distinct blunt tubercles (Page 1982), lopho-reticulate (with projecting anastomosing 
ridges forming an open angular reticulum) (Harris 1955), having a coarse network, 
tubercular in outline (Verma 1958), verrucate, appearing reticulate (Pant & Misra 
1976), reticulate with prominent lamellae (Knox 1951), reticulate with muri which 
vary remarkably in height and width (Ferrarini et al. 1 986), and scabrose-foveolate or 
baculate (Moe 1974). 0. lusitanicum spores, on the other hand, have been commonly 
described as smooth (Durieu de Maisonneuve 1857, Rothmaler 1964, Page 1982), 
though Knox (1 951 ) described them as irregularly pitted, Fernandes & Queirds (1 980) 
as finely reticulate with small areoles, Ferrarini et al. (1986)asfoveolate, andWildpret 
de la Torre et al. (1 974) as granular. Spores of 0. azoricum have been referred to as 
tuberculate (Rothmaler 1964), coarsely reticulate, not tuberculate (Bailey 1880) and 
reticulate-verrucate (Fernandes & Queirds 1980). 

There is obviously a lack of uniformity of terminology throughout the literature, 
although there is considerable variation in spore wall ornamentation between spores 
of different populations of a species, and even of one individual. Spores of 0. 
lusitanicum were more uniform than those of the other two species. SEM studies 
revealed 0. lusitanicum spores were irregularly pitted with many small, irregularly 
shaped pits, these being more numerous on the distal surface (Fig. 5e, f). These pits are 
visible in LM, and the outline of spores in equatorial view appear almost smooth, but 
slightly crenate. 

O. vulgatum spores are much more variable than those of 0. lusitanicum. The 
majority of spores observed under SEM were deeply pitted, especially on the distal 
surface where the many large, irregularly shaped pits were crowded together, the pit 
walls thus forming a very irregular reticulum (Fig. 5b). Onthe proximal surfacethepits 
are usually smaller and further apart (Fig. 5a). Some spores studied, however, have 
more widely-spaced pits, still forming a reticulum, but with some walls much wider 
and flat-topped. This supports the view that the reticulum comprises pit walls rather 
than tubercles. Under LM the equatorial outline of all spores is irregularly tuberculate, 
but the reticulum on the distal surface is also visible. 

The spores of 0. azoricum are generally intermediate in ornamentation between 
the above species. Most spores studied by SEM were irregularly pitted on both 
surfaces, the irregularly shaped pits being generally further apart on the proximal 
surface, and slightly larger than the pits of 0. lusitanicum spores (Fig. 5c, d). LM 
observations also reveal these pits, and the equatorial view is not as smooth as in 0. 
lusitanicum spores, but usually more regular and not as tuberculate as in spores of O. 
vulgatum. However, some O. azoricum spores have larger pits which are crowded 
together, forming a reticulum as in O. vulgatum. 



182 



FERN GAZETTE: VOLUME 13 PART 3 (1987) 




FIGURE 5. SEM photographs of Ophioglossum spores. All x 1300. a, b, O. vulgatum. M. Bell, July 
1977, Aston Clinton, Buckinghamshire; a, proximal view and b, distal view. c,d, O. azoricum, F. 
Townsend, June 1863, St. Agnes, Scilly Isles; c, proximal view and d, distal view. e,f, O. 
lusitanicum, M. Dawber, 21 January 1886, Guernsey, Channel Islands; e, proximal view and f, 
distal view. All spores untreated. 



THE STATUS OF OPHIOGLOSSUM AZORICUM IN THE BRITISH ISLES 183 



Thus, three distinct types of spore have been found in British Ophioglossum 
corresponding to the three taxa and these are shown in Fig. 5. However, there is 
considerable variation and thus overlap, particularly between 0. azoricum and 0. 
vulgatum, which means spore ornamentation is not a reliable character in 
distinguishing these two species. The spore wall surface between the pits is granular 
in most samples of the three species, though in some specimens the granules appear 
to coalesce. It is difficult to be sure that spores of equal maturity are examined in every 
case, and it is likely that some of the differences in the spore wall ornamentation may 
be attributed to different degrees of maturity, as some wall layers are not deposited 
until just prior to sporangial dehiscence, and layers which have been laid down may 
be entirely or partly lost before spores are examined (Lugardon 1978). 

DISTRIBUTION AND ECOLOGY 
Distribution of Ophioglossum in Europe 

Durieu de Maisonneuve (1 857) discusses at some length what was probably the first 
discovery in Europe of the taxon now called 0. azoricum. It was found in two quite 
different French localities, Pocancy near Paris by Puel and Vigineix in 1 846, and in the 
coastal dunes at Cap-Ferret near Arcachon in 1 857 by Durieu de Maisonneuve. These 
discoveries caused great excitement among local botanists, who debated whether the 
plants were 0. /us/tan/cum, a reduced form of 0. vulgatum or indeed, a third species. 
The taxon was described and named a few years later by Cosson and Germain (1861) 
as 0. vulgatum var. ambiguum. 

0. azoricum has since been reported from a number of European countries, and 
the distribution given for this species in Atlas Florae Europaeae (Jalas & Suominen 
1 972) is western Europe, from Iceland in the north, Great Britain, Ireland and France, 
to the Azores, Portugal, Corsica and Sardinia in the south, and Czechoslovakia and 
Poland in central Europe. Other records give 0. azoricum in Spain (e.g. Casasecas etal. 
1982, Lopez Gonzalez 1983, Monge & Velayos 1984) and (as 0. vulgatum f. 
ambiguum) in Italy (Fiori 1943). Lobin (1986),. in his discussion of Ophioglossum in 
Macaronesia, reports 0. azoricum from Madeira as well as the Azores, but regards 
reports for the Canary Islands and Cape Verde Islands as erroneous. Many of these 
sites are coastal or at least have an atlantic climate; others are continental, which is 
interesting since specimens found inland in Britain with an atlantic climate would 
normally be considered to be small 0. vulgatum, and not candidates for 0. azoricum. 
However, since the identity of this taxon is probably as confused in Europe as it is in 
Britain, these records may not reflect its true distribution. 

0. lusitanicum is essentially a species of the Mediterranean region and western 
Europe, ranging from Portugal in the west to Turkey in the east. This is again a 
predominantly coastal plant with some inland stations. The third species, 0. vulgatum, 
on the other hand, occurs throughout most of Europe, but is rarely found in the 
Mediterranean region. 

Distribution and ecology of Ophioglossum in Britain 

Ophioglossum vulgatum is relatively common throughout Britain, particularly in 
lowland central and south-eastern England. It is less frequent in the upland areas of 
Scotland, Wales and south-west England. Fronds generally appear in late April or May, 
mature in June or July and die down in August to September. 

This species grows in a wide variety of habitats. Most commonly it may be found in 
old damp meadow pastures, but it also occurs on the drier chalk downlands of 
southern England, as well as in old chalk pits and marl pits. Damp, often peaty 
depressions in heathland provide a suitable habitat, as do sand-dune slacks. In E. 
Norfolk 0. vulgatum grows in mown Phragmites mire on shallow peat over lake muds, 



184 FERN GAZETTE: VOLUME 13 PART 3 (1987) 



and the species has also been collected from the edge of a wet willow wood where it 
was growing in shallow water on flint shingle. Colonies are oftenfoundon sandy soils 
growing in association with bracken, and also in woods, copses and hedgebanks, 
sometimes in deep shade. 

By contrast, 0. lusitanicum is a very local species, found only in a few sites on St. 
Agnes in the Scilly Isles, and Guernsey in the Channel Islands. Plants may be evident 
above ground as early as September, maturing from November to March and dying 
down bytheendof April. Timing depends on the seasons, which may vary considerably 
from year to year. This tiny fern is mainly to be found in short turf on the moist peaty or 
sandy soil of coastal rocky downs and cliff-tops. These sites usually face south or 
south-west and are relatively exposed. 

Jermy et al. (1978) record the distribution of O. azoricum as Channel Islands, 
Scilly Isles, Lundy (Devon), Studland (Dorset), Skomer Island (Pembrokeshire), 
Ravenglass (Cumberland), Inner Fame (Northumberland), St. Kilda (Outer Hebrides), 
Dunnett Head (Caithness), and a number of sites in Merioneth, Caernarvon, Anglesey, 
Shetland Islands, Orkney Islands and the west coast of Ireland (Donegal, Sligo, Mayo 
and Kerry). These records were based on data culled from herbarium specimens, floras 
and field records, using the criteria of sporangial number (14 or fewer pairs), sterile 
blade size (less than 3.5cm long) and habitat (occurring in short turf near the sea). 
There are a number of specimens from other sites in various British herbaria labelled 
0. azoricum. However, I am doubtful that many of the specimens from these and well- 
known localities are anything other than small forms of 0. vulgatum, and have seen 
more convincing specimens from inland sites in Hampshire. O. azoricum follows the 
same seasonal growth pattern as O. vulgatum, being evident from April to September 
and maturing in June or July. It is reported to grow in short, grazed turf on top of cliffs 
and sea crags (e.g. in Orkney), and also in sand dune slacks (e.g. at Ravenglass). These 
are generally damp, often exposed sites, similar to those preferred by O. lusitanicum. 

I have seen the following specimens which are distinct in habit from O. vulgatum 

and have small fronds and relatively few sporangia thereby falling within the present 

concept of O. azoricum: 

V.-c. O, Channel Islands (Guernsey, L'Ancresse Common: M. Dawber, 14 June 1884 (BM, LIV), 
June 1886 (DBN, NMW), 18 June 1886 (BM), 14June 1887 (CGE), 28 May 1888 (OXF), 27 May 
1890 (BM), 29 May 1890 (DBN); J.D. Gray, June 1894 (BM, CGE); W.W. Reeves, June 1885 
(OXF)). V.-c. 1, Scilly Isles (W. Curnow, s/nerf/e (E). St. Agnes: J. Parslow, 15 May 1980(BM); F. 
Townsend, June 1863 (BM). St. Martins: J. Cunnack, June 1877 (CGE), June 1881 (BM); W. 
Curnow, June 1 877 (BM), 4 June 1 877 (BM), July 1 878 (BM, CGE, E, OXF); J. Ralfs, June 1 877 
(OXF, RNG),4July 1 877 (BM, CGE, E, K); B.V. Tellam, 1 877 (BM), 3 July 1877 (BM); F. Townsend, 
June 1862 (CGE)). V.-c. 4, North Devon (Lundy: A. Cleave, June 1982 (BM)). V.-c. 11, South 
Hants (near Lyndhurst: J.M. Camus & A.M. Paul, 2 July 1 985 (BM)). V.-c. 48, Merioneth (Morfa 
Harlech: DA. Jones, 1 July 1 898 (BM)). V.-c. 49, Caernarvon (Morfa Dinlle: R.H. Roberts, June 
1 961 (NMW)). V.-c. 52, Anglesey (Newborough Warren: R.H. Roberts, July 1 956 (NMW)). V.-c. 
70, Cumberland (Ravenglass: T. Dargie, Aug. 1971 (BM); A.C. Jermy & A.M. Paul, 14826, 26 
Sept. 1 980 (BM)). V.-c. 1 1 0, Outer Hebrides (St. Kilda: R.M. Barrington, June 1 881 (BM). Scarp: 
W.S. Duncan, 26 Sept. 1 890 (BM), summer 1 892 (BM), July 1 892 (BM). South Uist: A. Cleave, 
June 1 983 (BM)). V.-c. 111. Orkney (Calf of Flotta: J.T. Boswell, Aug. 1 880(OXF); W.I. Fortescue, 
July 1 878 (BM); H.H. Johnston, 2 Aug. 1 878 (BM, E). Eday: H.H. Johnston, 1 2 July 1 883 (BM). 
Fara: J.T. Boswell, Aug. 1 880 (BM); H.H. Johnston, 20 Aug. 1 880 (BM). Orphir: H.H. Johnston, 9 
Sept. 1 880 (OXF), 1 5 Aug. 1 881 (BM). Swanbister: J. Boswell Syme, Aug. 1 873 (BM), J.T. Syme, 
Sept. 1 855 (BM)). V.-c. 1 1 2, Shetland (Northmaven, Brei Wick: W.H. Beeby, 6 June 1 896 (BM, K, 
LIV). Papa Stour: W. Scott & R.C. Palmer, 19 July 1959 (OXF)). V.-c. H1, South Kerry (Great 
Blasket: R. Lloyd Praeger, June 1912 (BM, DBN). Three Sisters Head: R.W. Scully, 3083, Julv 
1885 (DBN)). 

DISCUSSION 
Species delimitation in Ophioglossum is a worldwide problem. These simple plants 
have few characters relative to other ferns, and classifications have often been based 
on features which in other pteridophytes would be considered unnoteworthy. Of the 



THE STATUS OF OPHIOGLOSSUM AZORICUM IN THE BRITISH ISLES 185 



three species recognised in Britain, 0. lusitanicum is clearly distinct in terms of 
growing season, size, frond shape, venation, epidermal cells and stomata, and spore 
ornamentation. The other two taxa, however, were found to overlap in all these 
respects to such an extent that many herbarium specimens could not be determined 
with any degree of certainty. The above features do not really help to separate O. 
azoricum from small specimens of O. vu/gatum, though it is perhaps possible that 
more extensive studies might reveal significant differences between the taxa that 
were not evident in this study. 

It is common for juvenile plants, especially ferns, to be fertile despite their small 
size and with a morphology often quite different from that of mature specimens. There 
is no reason why this should not apply to Ophioglossum, and it may be that sorrfe plants 
named 0. azoricum are merely juvenile forms of O. vu/gatum. 

Wagner et a/. (1 981 ) point out that those familiar with Ophioglossum in the field 
are more likely to have reliable taxonomic opinions than herbarium workers. It is true 
that different populations undoubtedly appear very distinct in the field, but how much 
of this is due to ecotypic variation? Wagner et a/. (1981) showed experimentally the 
range of variation in frond size and venation in a single species. 

It seems quite possible that O. vu/gatum would not grow as luxuriantly in exposed 
or heavily grazed coastaj sites, as in a damp meadow, just as other pteridophyte and 
spermatophyte species growing in adverse conditions are stunted and different in 
habit to specimens in more favourable situations. An example of this may be the 
Hampshire specimens referred to above which grow in small damp depressions in 
areas regularly trampled by humans and grazed by rabbits. These plants greatly 
resemble coastal specimens by 0. azoricum in size and habit. Are the Hampshire 
specimens true O. azoricum? Alternatively, are many, if not all the populations hitherto 
called 0. azoricum in reality just an ecotype of O. vu/gatum? \X is interesting to note a 
comment by P. Taylor (unpubl. data) that he has produced plants similar to those of 0. 
azoricum on Skomer Island by starving ordinary 0. vu/gatum. Miss E. Bullard (pers. 
comm. via A.C. Jermy) likewise considers that one would expect frond size and 
sporangial number to be reduced in the impoverished soils of Orkney where 0. 
azoricum is found. I hope to cultivate under uniform conditions plants which appear 
distinct from different habitats, and also to take plants from a single population and 
grow them under a variety of conditions, to see if differences are maintained inthefirst 
instance or generated in the second. 

The angle at which the sterile blade is held was noted to be a useful field 
character. Specimens of 0. azoricum from Ravenglass and Lundy had distinctly 
reflexed fronds compared to the upright habit of 0. vu/gatum. This feature is, however, 
not clearly shown by herbarium specimens. There is an interesting series of 
collections from Swanbister, Orkney, made by J.T. Boswell Syme over a period of 20 
years. The samples maybe from more than one population, butthere is a great range of 
variation in size. In general, the larger the specimen, the more numerous the 
sporangia. One sheet in particular (BM) has many specimens collected in 1 855 with 
two fronds, sterile blades up to 50-60mm long, 20-25mm wide and up to 19 pairs of 
sporangia. Many of these do have distinctly reflexed fronds. Most of the other 
collections from 1855, 1873 and 1875 comprise smaller individuals with 7-1 6(-1 8) 
sporangial pairs and sterile blades 7-35 x 4-1 5mm. A high proportion of these have 
more than one frond per plant and sterile blades reflexed to some degree. Plants in the 
first-mentioned collection, although larger than O. azoricum is usually defined, do 
appear to have the characteristic habit of this taxon. Even this feature may be the result 
of the habitat; in the taller vegetation of 'vu/gatum' sites fronds are more likely to be 
erect and tall, whereas in the short turf of 'azoricum' sites they are more likely to be 



186 p ERN GAZETTE: VOLUME 1 3 PART 3 (1 987) 



small and reflexed. Ecological data of any sort is often absent from herbarium 
specimen labels, so extensive field studies would be most useful, collecting data on 
frond size and habit, ecology including observations on associated species, height of 
vegetation etc. Photographs to accompany herbarium specimens would contribute 
valuable information. 

In conclusion, it is evident that although there are some specimens which are 
distinct in size and habit from typical O. vulgatum and which may be confidently 
termed 'O. azoricum', there are many individuals and populations intermediate 
between these two extremes. This prevents satisfactory separation of the two species 
on morphological grounds. They do not seem to be as distinct in measurable terms as 
the two North American varieties of 0. vulgatum (now recognised as species) studied 
by Wagner (1971). Although in many respects it seems that O. azoricum should be 
synonymised with 0. vulgatum, the fact that some populations are so manifestly 
distinct with their tiny reflexed leaves leads me to accept it is as a separate species for 
the time being pending further research. 

I did not find substantial evidence of O. azoricum being intermediate between the 
other two British taxa, but rather found it to be merely at one end of the range of 
variation exhibited by 0. vulgatum under differing conditions. This factor, coupled with 
the distinctness in many ways of 0. lusitanicum from 0. azoricum, offers little 
morphological support to the proposal of the latter's hybrid origin. 

As well as extensive field studies, a cytological survey of different populations of 
Ophioglossum (including those in the Azores) is necessary to help establish the status 
of 0. azoricum in Britain and, indeed, Europe. If there are three cytologically different 
taxa involved, chromosome counts combined with morphology may enable the two 
problem taxa to be more satisfactorily defined. 

ACKNOWLEDGEMENTS 
Thanks are due to the staff of the following herbaria for lending specimens: CGE, DBN, 
E, K, LIV, NMW, OXF, RNG. I am grateful to Miss J.M. Camus, Dr M. Gibby, Mr A.C. 
Jermy and Mr W. Lobin for helpful discussion. I would also like to thank Mr A. Cleave 
and Mr K.H. Hyatt for allowing me to use their photographs. 

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188 FERN GAZETTE: VOLUME 13 PART 3 (1987) 

SHORT NOTE 

ADIANTUMxSPURIUM, A NEW NAME FOR A. x VILLOSOLUCIDUM 

In a series of papers entitled "Cytotaxonomic studies of the ferns of Trinidad" (Jermy 
and T. Walker, 1985, in Bull. Brit Mus. (Nat. Hist.), Botany ser. 13 (2): 133-276) the 
name Adiantum x villosolucidum Jermy and T. Walker, was used as an epithet for the 
hybrid between A lucidum (Cav.) Swartz and A villosum L. This hybrid was found on 
several occasions in Trinidad where the parents grew together, and is most likely to be 
found elsewhere throughout the sympatric range ofthe two species inthe West Indies 
and South America. 

It has been pointed out that according to Article H. 1 0.3 in the International Code of 
Botanical Nomenclature (Voss et af., 1983, Regnum Vegetabile 111, Utrecht) 
designations consisting of the names of the parents combined with only the 
termination of one epithet changed is considered to be a formula and not a true epithet. 
The name villosolucidum is therefore invalid and to rectify the mistake we propose 
here the following substitute: 

Adiantum x spurium Jermy and T. Walker, nom. nov. based on Adiantum x 
villosolucidum Jermy and T. Walker, \r\Bull. Brit. Mus. (Nat. Hist), Botany ser. 13 (2): 
256 (1985). Typus: Trinidad, Tacarigua Ward, Tunapuna, 2 mile along Caura Road, c. 
60m alt., on shady bank, 3 April 1966, M.G. & T.G. Walker T10588 (holotype, BM: 
isotype herb. Walker). Paratypes: Same locality, date and collectors, T10589 (TRIN), 
T10590 (NY), T10591 (CR) and T10592 (BM). 

A.C. JERMY 
British Museum (Natural History), Cromwell Road, London, SW7 5BD 

T.G. WALKER 
Department of Plant Biology, University of Newcastle upon Tyne, NE1 7RU 



FERN GAZ. 13(3) 1987 189 

OBITUARY 

CHING REN CHANG 1898-1986 

Ching Ren Chang was born on 16 January, 1898, in Wujing County of Kiangsu 
Province. His first academic course was on forestry and he graduated from Kingling 
University, Nanking in 1 925. For the next three years he lectured at the Department of 
Biology, Southeast University, Nanking. Between 1 927 and 1 932, Ching was Head of 
the Botany Section of the Metropolitan Museum, Nanking, moving to Peking, as Head 
of the Herbarium of the Fan Memorial Institute of Biology, in 1932 until 1945. 

During these early years Ching Ren Chang started to specialise on ferns and 
visited the Herbarium of the Hong Kong Botanical Garden in 1 926 to study the fern 
collections there. He followed this up with a major European visit 1929-32 during 
which time he studied under Carl Christensen in Copenhagen, and visited the British 
Museum and Kew and herbaria in Austria, Czechoslovakia, France, Germany and the 
Netherlands. In all of these he photographed as many fern types as he could find and 
this collection, now in the National Herbarium, Beijing, is an important reference 
database. 

Between 1 945 and 1 955 his energies were directed to agricultural improvement, 
but on the setting up of the Academia Sinica in 1955 he was made a Member, and 
Director of the Phytotaxonomy Department of the Botanical Institute in Beijing. He 
continued working on ferns to the end, latterly as an Adviser of the Botanical Institute, 
publishing over 140 papers and books. A full list will be published in the Bulletin of 
Pteridology in Taxon. He has trained and inspired many young Chinese botanists and 
leaves a strong contingent of pteridologists in both Academia and University 
Departments. 

Ching Ren Chang married Zuo Jin Fu in 1933, who passed away in 1964. He 
leaves a son, Ching Li-Ming, with whose family he lived during his last years. 

A.C. Jermy & K.H. Shing 



190 FERN GAZETTE: VOLUME 13 PART 3 (1987) 

BRITISH PTERIDOPHYTE RECORDS 

Compiled by A.J. Worland 

The records are presented thus: 100km square/1 Okm square followed by the 
recorder's name. Listed additions have been received up to the end of March 1987. 

POST 1980 

1.2 Lycopodium clavatum 17/52, 17/64 J. Clark. 34/19, 35/40, 35/41, 35/52 
G. Halliday. 

3.1 Huperzia selago 34/19, 34/28, 34/39, 35/60, 35/70, 35/71, 35/81 G. Halliday 

4.1 Diphasiastrum alpinum 17/52 J. Clark 

6.2 Isoetes echinospora 07/94, 17/04, 17/26 A.C. Jermy & J. Clark 
7.1 Equisetum hyemale 17/45 J. Clark 

7.4 Equisetum arvense 17/05 A. Stirling 

7.7 Equisetum palustre 17/05 A. Stirling 

8.1 Botrychium lunaria 17/22, 17/31 J. Clark. 34/28, 35/23, 35/70, 35/80, 35/81 

G. Halliday 

9.1 Ophioglossum vulgatum 17/41 J. Clark 

11.1 Cryptogramma crispa 17/42 J. Clark 

14.1 HymenophyNum tunbrigense 20/88 M.H. Rickard 

14.2 HymenophyNum wilsonii 35/34, 35/53 G. Halliday 
16.1-3 Polypodium vulgare agg. 17/05 A. Stirling 

16.1 Polypodium vulgare 17/25 U.K. Duncan 

16.2 Polypodium interjectum 07/93, 07/94, 17/04, 17/05 A. Stirling 
17/15, 17/25, 17/26 J. Clark 

17.1 Pteridium aquilinum 07/93, 17/05 A. Stirling 

20.1 Oreopteris limbosperma 07/94 U.K. Duncan 

21.1 Asplenium scolopendrium 17/04, 17/25 AC. Jermy & J. Clark 
35/14, 35/15, 35/25, 35/26, 35/36, 35/41, 35/43 G. Halliday 

21.2 Asplenium adiantum-nigrum 07/93, A.C. Jermy & J. Clark. 17/72, 17/74 J. Clark 

21.7 Asplenium trichomanes 07/93 A. Stirling. 35/36 G. Halliday 
21.7a Asplenium trichomanes subsp. trichomanes 17/72 J. Clark 

21.8 Asplenium viride 07/93 A. Stirling. 1 7/94 A.C. Jermy & J. Clark. 35/56 G. Halliday 
35/56 G. Halliday 

21.9 Asplenium ruta-muraria 17/43 J. Clark 

24.1 Cystopteris fragilis 17/72, 17/73 J. Clark. 34/19, 34/28, 35/14, 35/27, 35/36, 

35/44, 35/45 G. Halliday 

22.1 Athyrium filix-femina 07/93 A. Stirling 

26.2 Polystichum aculeatum 34/19, 34/28, 35/00, 35/35 G. Halliday 

27.2 Dryopteris filix-mas 07/93 A. Stirling 

27.3 Dryopteris affinis 17/04 A. Stirling. 17/26 U.K. Duncan 
27.9 Dryopteris dilatata 17/04 A.C. Jermy & J. Clark 

28. 1 Blechnum spicant 1 7/04 A.C. Jermy & J. Clark 

29.1 Pilularia globulifera 1 7/04 A.C. Jermy & J. Clark 

PRE 1980 (AND POST 1960 IN BRACKETS) 

3. 1 Huperzia selago (07/94), (1 7/04) U.K. Duncan. 17/15 A.C. Jermy & J. Clark. 

(17/25) V. Gordon. 17/26 J. Clark 
5.1 Selaginella selaginoides (07/94) U.K. Duncan. 1 7/04 A. Stirling 

17/15, 17/26 A.C. Jermy & J.Clark. 17/25 J. Clark 

7.3 Equisetum fluviatile 07/94, 1 7/04 A Stirling. 17/15, 17/25 J. Clark 
17/26 U.K. Duncan 

7.4 Equisetum arvense 07/94, 17/04 A. Stirling. 17/15, 17/25, 17/26 J. Clark 
7.0* Equisetum sylvaticum (17/25) V. Gordon 

7.7 Equisetum palustre 07/94, 1 7/04 A. Stirling. 17/15, 17/25, 17/26 J. Clark 

8.1 Botrychium lunaria 17/15 J. Clark 

9. 1 Ophioglossum vulgatum (17/15) U.K. Duncan. 1 7/25, 1 7/26 J. Clark 

10.1 Osmunda regalis 17/15 J. Clark. 17/25 A.C. Jermy & J. Clark. (17/26) U.K. Duncan 

14.2 Hymenophyllum wilsonii (17/15) U.K. Duncan. 17/26 J. Clark 

16.1-3 Polypodium vulgare agg. 07/93, 07/94, 17/04 A. Stirling. 17/15, 17/25, 17/26 

J. Clark 
1 7. 1 Pteridium aquilinum 07/94, 1 7/04 A. Stirling. 17/15, 1 7/25, 1 7/26 J. Clark 

21.1 Asplenium scolopendrium (07/94), (17/15), (17/26) U.K. Duncan 

21.2 Asplenium adiantum-nigrum 07/94 A. Stirling. 17/04 A C. Jermy & J. Clark. 
17/26, 17/33, 17/34, 17/43, 17/44 J. Clark 

21.6 Asplenium marinum 07/94 A.C. Jermy & J. Clark. 17/15, 1 7/25 J. Clark. 

(17/26) U.K. Duncan 



BRITISH PTERIDOPHYTE RECORDS 191 



21.7 Asplenium trichomanes 1 7/04 A.C. Jermy & J. Clark. 1 7/1 5 J. Clark 

21.9 Asplenium ruta-muraria 17/15, 17/43 J. Clark 

21.10 Asplenium septentrionale 20/88 M.H. Rickard 

22.1 Athyrium filix-femina 07/94, 17/04 A. Stirling. 17/15, 17/25, 17/26 J. Clark 

24.1 Cystopteris fragilis 35/26, 35/34 G. Halliday 

27.2 Dryopteris filix-mas 07/94, 17/04 A. Stirling. 17/15. 17/25 J. Clark 
(17/26) U.K. Duncan 

27.5 Dryopteris aemula 17/15, 17/25 A.C. Jermy & J. Clark 

27.8 Dryopteris carthusiana 44/37 J. Bouckley 

27.9 Dryopteris dilatata 07/94 A. Stirling. (17/15) V. Gordon. 17/25 J. Clark. 
(17/26) U.K. Duncan 

28.1 Blechnum spicant 07/94 A. Stirling. 17/15, 17/25, 17/26 J. Clark 

RECORDS 
Would members and leaders of field meetings kindly send any new or updated records 
to the pteridophyte records recorder for checking and inclusion in the annual lists of 
updates and amendments. 



192 FERN GAZETTE: VOLUME 13 PART 3 (1987) 



THE FERN GAZETTE 

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The British Pteridological Society 
THE FERN GAZETTE 



VOLUME 13 PART 3 



1987 



CONTENTS 



Page 
MAIN ARTICLES 
T. Reichstein: a personal appraisal 

- Irene Manton 1 3 1 
Asplenium reichsteinii (- Asplenium fontanum x A. major/cum; Aspleniaceae: 
Pteridophyta), a new endemic fern hybrid from Mallorca, Balearic Islands 

- H. Wi/fried Bennert, Helga Rasbach & Kurt Rasbach 1 3: 

Notes about Asplenium I. Asplenium quezelii, a pseudo-endemic species 
identical with A. daghestanicum (Aspleniaceae: Pteridophyta) 

- R.L.L Viane 14: 

Hybridisation experiments between Asplenium see/osii and A. ce/tibericum 
(= A. see/osii subsp. g/abrum) (Aspleniaceae: Pteridophyta) 

-J.D.Lovis 151 

Observations of progeny of Athyrium filix-femina (Athyriaceae: 
Pteridophyta) from breeding experiments 

- J.J. Sen ne Her 15^ 

Asplenium tadei (Aspleniaceae: Pteridophyta), a new species from Turkey 

- C.R. Fraser-Jenkins & J.J. Schneller 163 

The fern herbarium of Col. F.J. Hutchison 

- John Edmondson 1 69 

The status of Ophioglossum azoricum (Ophioglossaceae: Pteridophyta) 
in the British Isles 

-A.M. Paul 173 



SHORT NOTE 

Adiantum x spurium, a new name for A. x villosolucidum 

- A.C. Jermy & T.G. Walker 

OBITUARY: CHING REN CHANG 
-A.C. Jermy & K.H. Shing 



188 
189 



REVIEWS 

•BRITISH PTERIDOPHYTE RECORDS 



142, 150, 162 
190 



(THE FERN GAZETTE Volume 13 Part 2 was published on 29th September 1986) 



Published by THE BRITISH PTERIDOLOGICAL SOCIETY, c/o Department of 
Botany, British Museum (Natural History), London SW7 5BD 



ISSN 0308—0838 

Metloc Printers Ltd., Old Station Road, Loughton, Essex 




IL 



Edited by 

J.A£rabbe 

& B.S.Parris 



FHE 
3R1T1SH 

PTER1D0L0G1CAL 
SOCIETY 



Volume 13 Part 4 



JS 



1988 



THE BRITISH PTERIDOLOGICAL SOCIETY 
Officers and Committee for 1 988 

President: Dr B.A. Thomas 
President Emeritus: J.W. Dyce 

Vice-Presidents: Dr R.E. Holttum, F. Jackson, A.C. Jermy, R. Kaye, G. Tonge 

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Treasurer: Dr N.J. Hards, 

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The Editor of the Fern Gazette, British Pteridological Society, 

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A.M. Paul, P. Ripley, Dr T.G. Walker, Dr A. Willmot 

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FERNGAZ. 13(4)1988 193 

LIB 

A CHROMOSOME COUNT FROM AZOLLA FILICULOIDES 
(AZOLLACEAE: PTERIDOPHYTA)* 

YOU XING LIN** and ANNE SLEEP 
Department of Pure and Applied Biology, The University, Leeds, LS2 9JT, 

England 

ABSTRACT 
Azolla filiculoides has been investigated cytologically for the first time. Counts from 
microsporangia establish its chromosome number as n = 22. 

INTRODUCTION 
To date there have been very few cytological investigations of Azolla (the Water Fern). 
The minute chromosomes present technical problems which are compounded by the 
delicate nature of the roots and a scarcity of fertile material at the right stage. The 
extremely small chromosome size also causes considerable difficulty in the 
establishment of unequivocal counts. The cytology of Azolla has thus lagged behind 
research into other aspects of its biology. Recently much interest has focussed on 
this genus because of its nitrogen-fixing ability; this arises from the symbiotic 
relationship between Azolla and the endophytic cyanobacterium (or blue-green alga), 
Anabaena azollae, which is invariably present in the air chambers in the upper lobes 
of its leaves. The consequent economic importance of Azolla has long been recognized 
in the Far East, where, particularly in China and Vietnam, A. pinnata R.Br, has been 
used for centuries as a green manure for increasing the fertility of paddy fields (Lumpkin 
1 985). During the last ten years various other species of Azolla have been introduced 
into China for the purposes of breeding and other research, and the arrival in Leeds 
of Y. X. Lin presented an opportunity to count one of the most vigorous of these 
introduced Azolla species, a particularly strong-growing form identified as A. filiculoides 
Lam. Azolla shows much variation in the morphology of its vegetative parts, and 
the same species can look quite different when growing under different environmental 
conditions. This phenotypic plasticity has led to problems over identification; the 
boundaries between species are unclear and in general the taxonomy of the genus 
is confused. 

MATERIAL 
The material used in this study came originally from the Forstbotanischer Garten 
Eberswalde der Humboldt, at the University of Berlin in East Germany. This culture 
was sent to the botanic garden of the Botanical Institute, Chinese Academy of Sciences, 
Beijing, in 1978, and was established in cultivation there. In 1986 material of the 
same collection was successfully brought into cultivation at Leeds. Sporangia were 
produced in the autumn of 1986 and again in 1987. Material of botanic-garden origin 
may sometimes be misidentified, especially in view of the problems outlined above. 
It was therefore thought desirable to check the identification of our culture as A. 
filiculoides, and to this end various morphological features were studied. Characters 
for the identification of A. filiculoides are summarized by Ashton & Walmsley (1984) 
and by Ruiz de Clavijo et al. (1 984). Fertile material of mature, well-grown A. filiculoides, 
one of two American species introduced and now widespread in Europe, is said to 
be very distinctive (the other, A. caroliniana Willd., is much more difficult to identify 
positively and its taxonomic status may be doubtful, see Derrick, Jermy & Paul 1987) 

* Dedicated to Prof. T. Reichstein on the occasion of his 90th birthday (20.7.1987) in recognition 
of his friendship and encouragement. 

**0n leave from the Institute of Botany, Chinese Academy of Sciences, Beijing, People's Republic 
of China. 



194 



FERN GAZETTE: VOLUME 1 3 PART 4 (1 988) 




PLATE 1 

I. Morphology of our cytologically attested Azolla material (A. filiculoides) : SEM photographs 

of leaves. 

Fig. 1. Upper epidermis showing the wide margin of 3-4 rows of large, hyaline cells 

and the central lamina bearing stomata and unicellular trichomes or papillae. 

Fig. 2. Enlarged view of the unicellular trichomes. 



AZOLLA FILICULOIDES 195 



Our material is a strong-growing plant, consisting, when mature, of both a 
decumbent, free-floating system as well as erect branches which can extend out 
of the water by as much as 5cm. When the emergent branches are well-developed 
the horizontal system tends to die back, forming a tangled brown mat on the water 
surface. The ascending growth habit arises particularly when the plant is growing 
in a limited space or under dry conditions. Lin (1980) has observed that on these 
upright branches the colourless and usually submerged lower lobes of the leaves 
also become green and develop /Ana^aena-containing cavities. The flattened, leafy 
stems of the horizontal system are elongate, and the branches distant one from another. 
In young plants the branching is dichotomous; on adult material unequal development 
at branch-points leads to a false dichotomy and branching of apparently irregular 
arrangement. The tips of the emergent branches tend to roll inwards to give a distinctive 
crisped appearance mentioned by many authors. Roots arise in no set pattern or 
position but acropetally at intervals along the underside of the floating stems. The 
ovate, imbricate, closely appressed branch-leaves agree with the description of Svenson 
(1944); they are alternate, greyish-green in colour, with subacute to obtuse tips and 
a broad, membranous margin (as described by Lawalree in Tutin 1964) formed of 
2-3 rows of large, hyaline cells (see Figs. 1 & 2). Unicellular trichomes (Lumpkin 
& Plucknett 1980, Pieterse, de Lange & van Vliet 1977) are borne on the surface 
of the epidermis (see Figs. 1 & 2). Sporocarps develop, usually in pairs, in the axils 
of leaves close to the branch apices. Either two microsporocarps (very rarely, two 
megasporocarps) or a microsporocarp and accompanying megasporocarp occur 
together. Within each microsporocarp there are numerous microsporangia, each 
containing 4-6 massulae. The massulae bear barbed glochidia which are clearly 
aseptate (see Fig. 3). Fowler and Stennett-Willson (1978) attach great importance 
to the structure of the megaspore wall in classification; although not studied in detail 
the megaspore sculpturing of our material agrees well with that illustrated by them. 
On the basis of these observations we have no hesitation in confirming the identification 
of our plants as A. filiculoides Lam. 

Voucher herbarium specimens of our material will be deposited in the national 
herbaria at Kew and the British Museum (Natural History), as well as in the herbaria 
of the Botanical Institute, Chinese Academy of Sciences, Beijing and of the International 
Rice Research Institute, Manila, Philippines. 

METHODS 
Young microsporangia were fixed in a solution of 3 parts absolute alcohol: 1 part 
glacial acetic acid, stained in acetocarmine and squashed according to the technique 
described by Manton (1950). Preparations were made permanent by McClintock's 
method (1 929). Suitable cells were photographed in phase-contrast on a Zeiss Ultraphot 
II microscope. Stereoscan photographs of leaves and glochidia were taken from material 
fixed in 1% osmium tetroxide, dehydrated through an ascending acetone series and 
critical-point dried before mounting and coating with gold. The specimens were 
examined on a CamScan Series III scanning electron microscope. 

CYTOLOGY 
Several cells in metaphase I yielded counts of n = 22 bivalents. The cell illustrated 
in Fig. 4 (with accompanying explanatory diagram in Fig. 5) is at late metaphase 
and some of the bivalents are already beginning to separate prior to anaphase I. 
One prominent pair is very much larger than the rest. Root-tip preparations by ourselves 
from the same material gave counts of 2n = c.44, which are in accordance with 
the meiotic observations. 



196 



FERN GAZETTE: VOLUME 1 3 PART 4 (1 988) 










/ 



! 



I I5(J 



J5p 



PLATE 2 

Fig. 3. SEM photograph of surface of massula showing the aseptate glochidia. 
II. Cytology: meiosis from microsporangium of A. filiculoides. 

Fig. 4. Late metaphase I showing 22 bivalents. One chromosome (arrowed) is much larger 

than the rest. Some bivalents are already beginning to separate. Magnification: x 2,400. 

Fig. 5. Explanatory diagram to fig. 4. n = 22. Magnification: x 3,800. 



AZOLLA FILICULOIDES 197 



DISCUSSION 

Love, Love and Pichi-Sermolli (1977) give base numbers for Azolla of x = 11, x = 
12. The latter number is based on two early reports, the first by de Litardiere (1921), 
who obtained a count of 2n = 48 from material of A. caroliniana. He used sectioned 
material, from which it is not always easy to establish a precise count. However, 
Tschermak-Woess and Dole^al-Janisch (1 959), in a paper on the cytology of Cyrtomium 
falcatum, state that they found 2n = 48 in material of A. caroliniana; this confirms 
the observation of de Litardiere. Tschermak-Woess and Dolezal-Janisch made counts 
from aceto-carmine squash preparations but they did not, unfortunately, illustrate 
any cells which would have confirmed their stated finding of 2n = 48. In view of 
our own report of n = 22 from A. filiculoides it is highly desirable that plants referred 
to A. caroliniana should be investigated cytologically. 

The practice of Love et al. (1977) in deducing base numbers on purely arithmetical 
grounds is unsound and has already been criticized elsewhere (Walker 1977). We 
follow Manton & Vida (1968 p. 365), quoted by Lovis (1977), in believing that the 
number used as a monoploid in any genus should be the lowest gametic number 
for which there is direct evidence. Since there is currently no evidence to the contrary, 
we take x = 22, and possibly also x = 24, to be the base numbers in the genus 
Azolla. The number x = 22 is also basic to the Osmundaceae. 

Our record agrees with the counts of n = 22 and 2n = 44 obtained by Loyal 
from both meiotic (1958) and mitotic (1975) preparations of A. pinnata from the Punjab. 
This species, like our material of A. filiculoides, may also be regarded as diploid. 
It is interesting that counts of n= 22 have now been recorded from each of the two 
sections (or subgenera) into which Azolla is divided: A. pinnata (together with A. 
nilotica Decne) is classified under section Rhizosperma, while A. filiculoides is placed 
in section Azolla (which also includes A. caroliniana, A. mexicana Presl, A. microphylla 
Kaulf. and A. rubra R.Br.). 

The chromosomes of Azolla are very small in size, almost at the limit of resolution 
of the light microscope, and are, according to Loyal (1958, 1975), the smallest 
chromosomes yet recorded in the Filicales. Loyal (1975) suggests that this represents 
an extreme phylogenetic reduction correlated with the aquatic habit of Azolla. In an 
interesting discussion in the same paper it is also suggested that the asymmetrical 
nature of the karyotype (we were able to confirm Loyal's observation that the largest 
chromosomes are approximately double the size of the smallest ones) is linked to 
the high degree of morphological specialization shown by Azolla in its reproductive 
structures and its symbiotic association with Anabaena azollae. 

Polyploidy is a common phenomenon in the homosporous ferns but, as pointed 
out by Klekowski & Baker (1 966), it does not appear to have developed to any appreciable 
extent in the heterosporous Pteridophyta (Selaginellaceae, Isoetaceae, Marsileaceae 
and Salviniaceae), where the cytological counts available so far reveal generally low 
numbers and remarkable uniformity. Many more counts have been made in the 
homosporous ferns and, as further material is studied, the presence of polyploidy 
in the heterosporous Pteridophyta, recently confirmed in Selaginella (M. Gibby pers. 
comm.), may prove to be more widespread. In this connection, it is interesting to 
note that in the genus Salvinia a pentaploid hybrid has been recorded by Tatuno 
& Takei (1969). Only further chromosome counts will settle the question of the base 
number of Azolla, and will reveal if any polyploid strains exist. To this end various 
species and strains of Azolla have already been introduced into cultivation at Leeds 
for further study. In the meantime it is hoped that this preliminary report and discussion 
of the literature will stimulate research into different aspects of the cytology of this 



1 98 FERN GAZETTE: VOLUME 1 3 PART 4 (1 988) 



tantalizingly difficult genus. 



ACKNOWLEDGEMENTS 

We are grateful to the Astbury Department of Biophysics in the University of Leeds 
for providing photomicrographic facilities, to Mr A. Holliday, photographer in the 
Department of Pure and Applied Biology, for his assistance, and to Mr A. Hick, of 
the same department, for the stereoscan photographs. Y. X. Lin also wishes to thank 
the Sino-British Fellowship Trust, the Universities' China Committee, the Great Britain- 
China Educational Trust and the British Council for their financial support. 



REFERENCES 

ASHTON, P.J. & WALMSLEY, R.D. 1984. The taxonomy and distribution of Azolla species in 

southern Africa. Bot. J .Linn. Soc. 89(3): 239-247. 
DERRICK, N.L, JERMY, A.C. & PAUL, A.M. 1987. Checklist of European Pteridophytes. 

Sommerfeltia 6: 1 -94. 
FOWLER, K. & STENNETT-WILLSON, J. 1978. Sporoderm architecture in modern Azolla. Fern 

Gaz. 77(67:405-412. 
KLEKOWSKI, E.J. & BAKER, H.G. 1 966. Evolutionary significance of polyploidy in the Pteridophyta. 

Science, N.Y. 753.305-307. 
LAWALREE, A. 1964. Azolla. In TUTIN, T.G. et al. Flora Europaea I: 25. Cambridge University 

Press. 
LIN, Y.X. 1980. A systematic study of the family Azollaceae with reference to the extending 

utilization of certain species in China. Acta Phytotax. Sin. 18(4): 450-456. (In Chinese, 

English summary.) 
LITARDIERE, R. de. 1921. Recherches sur I'element chromosomique dans la caryocinese 

somatique des Filicines. Cellule 31: 255-473. 
LOVIS, J.D. 1977. Evolutionary patterns and processes in ferns. In PRESTON, R.D. & 

WOOLHOUSE, H.W. Advances in Botanical Research 4: 229-41 5. Academic Press. 
LOVE, A., LOVE, D. & PICHI-SERMOLLI, R.E.G. 1977. Cytotaxonomical Atlas of the Pteridophyta. 

J. Kramer, Vaduz. 
LOYAL, D.S. 1 958. Cytology of two species of Salviniaceae. Curr. Sci. 27(9): 357-358. 
LOYAL, D.S. 1975. Chromosome size and structure in some heterosporous ferns with a bearing 

on evolutionary problems. In KACHROO, P. Advancing frontiers in cytogenetics: 293-298. 

Delhi. 
LUMPKIN, T.A. 1985. Advances in Chinese research on Azolla. Proc. R. Soc. Edinb. 86B: 161- 

167. 
LUMPKIN, T.A. & PLUCKNETT, D.L 1980. Azolla: botany, physiology and use as a green manure. 

Econ. Bot. 34: 1 1 1 -1 53. (Reprinted 1 982 by Westview Press.) 
MANTON, I. 1950. Problems of cytology and evolution in the Pteridophyta. Cambridge University 

Press. 
MANTON, I. & VIDA, G. 1968. Cytology of the fern flora of Tristan da Cunha. Proc. R. Soc. 

B. 7 70.361-379. 
McCLINTOCK, B. 1929. A method for making acetocarmine smears permanent. Stain Technol. 

4: 53-56. 
PIETERSE, A.H., DE LANGE, L. & VAN VLIET, J. P. 1977. A comparative study of Azolla in the 

Netherlands. Acta Bot. Neerl. 26(6): 433-449. 
RUIZ DE CLAVIJO, E., MUNOZ, J. & SALVO, A.E. 1984. Sobre la presencia de Azolla filiculoides 

Lam. en Espana. Acta Bot. Malacit. 9: 129-132. 
SVENSON, H.K. 1 944. The New World species of Azolla. Am. Fern J. 34(3): 69-84. 
TATUNO, S. & TAKEI, M. 1969. Cytological studies of Salviniaceae I. Karyotype of two species 

in the genus Salvinia. Bot. Mag., Tokyo 82: 403-408. 
TSCHERMAK-WOESS, E. & DOLEZAL-JANISCH, R. 1959. Uber die karyologische Anatomie 

einiger Pteridophyten sowie auffallende Unterschiede im Kernvolumen bei Cyrtomium 

falcatum. Ost. Bot. Z. 106: 315-324. 
WALKER, T.G. 1977. Review in Fern Gaz. 1 1(5): 329-330. 



FERN GAZ. 13(4)1988 199 



CYTOLOGICAL AND ANATOMICAL OBSERVATIONS ON 

TMESIPTERIS (TMESIPTERIDACEAE: PTERIDOPHYTA) 

SPECIES FROM NEW CALEDONIA 

A.F. BRAITHWAITE 
Botany Department, University Park, Nottingham, NG7 2RD 

ABSTRACT 

Cytological and anatomical observations on three of the four species of Tmesipteris 
recognised in New Caledonia are presented. The tree fern epiphytes, T. lanceolata 
Dang, and T. sigmatifolia Chinn., aretetraploid(n= 104) with sclerencnymatous pith 
cells in the aerial shoot and the terrestrial T. oblanceolata Copel. subsp. linearifolia 
subsp. nov. is octoploid (n = c.208) with a parenchymatous pith. No material of the 
fourth species, the terrestrial T. vieillardii Dang., was available during the present 
investigation but its stem anatomy showing a parenchymatous or collenchymatous 
pith is well documented in the literature. The chromosome numbers and pith types 
fall into the same patterns as those previously reported in the literature for other 
members of the genus from Australia, New Zealand, Solomon Islands and Vanuatu. 
From the evidence available at present for the genus as a whole, the chromosome 
number does not appear to be correlated with either pith type or ecology of the 
species. There is perhaps a closer, though not complete, correlation between pith 
type and ecology but the significance of this observation is at present not clear. 

INTRODUCTION 
The polyspecific nature of the genus Tmesipteris first put forward last century (Brown 
1806, Endlicher 1833, Dangeard 1890-91) is now widely accepted (Wakefield 1944, 
Barber 1 957, Chinnock 1 975, Braithwaite 1 973, 1 986). While much of the evidence 
comes from conventional morphological studies of herbarium specimens; cytological 
(Barber 1957, Braithwaite 1986), anatomical (Dangeard 1890-91, Sahni 1925, 
Braithwaite 1 973, 1 986) and field studies (Barber 1 957, Chinnock 1 975) have made a 
valuable contribution, particularly in Australia, New Zealand, Solomon Islands and 
Vanuatu. There are some anatomical observations available in the literature for the 
New Caledonian species (Dangeard I.e., Sahni I.e.) but up to now no information has 
been available on their chromosome numbers. The present paper records some 
cytological and further anatomical and ecological observations on the New Caledonian 
species. 

The majority of the earlier studies of Tmesipteris in New Caledonia placed the 
different forms under the type and original species T. tannensis (Spr.) Bernh. 
(Mettenius 1861, Diels 1906, Jeanpert 1911, 1912, Bonaparte 1921). The only 
exception was Dangeard (1890-91) who described one very distinctiveterrestrialform 
as T. vieillardii. At the same time he described T. elongatum and T. lanceolatum from 
Australia and recognised two earlier species, T. tannensis and T. truncatum R.Br., 
from Australia and New Zealand. Despite Dangeard's very detailed morphological and 
anatomical studies, it was not until the early 1920's that a polyspecific concept for 
Tmesipteris in New Caledonia was accepted. 

Compton (1922) recognised three species, namely T. lanceolata Dang., T. 
tannensis (Spr.) Bernh. and T. vieillardii Dang. Sahni (1925) examined Compton's 
material anatomically and maintained T. vieillardii, but on the basis of similarities in 
their anatomy reduced all the other forms to T. tannensis as var. lanceolata, var. typica 
(tannensis), which was a form of T. lanceolata, and var. elongata. The last epithet 
referred to the material placed under T. tannensis by Compton (1 922) and was for a 
time to become the name most commonly used in European herbaria for the species 
now known as T. sigmatifolia Chinn. 

Author's note added in proof: 

This paper was submitted in February 1 987 and type set for the 1 987 issue but space constraints 

prevented its publication in that issue. It does not, therefore, include the results subsequently 

reported by Patrick Brownsey and John Lovis in their excellent paper on the New Zealand species. 

See Brownsey, P.J. and Lovis, J.D. 1 987. Chromosome numbers for the New Zealand species of 

Psilotum and Tmesipteris, and the phylogenetic relationships of the Psilotales./V. Z. Jour. Bot. 25: 

439-454. 



200 FERN GAZETTE: VOLUME 1 3 PART 4 (1 988) 



Brownlie (1969) in the most recent fern flora of New Caledonia recognised only 
two species, T. lanceolata and T. vieillardii. The material originally referred to T. 
tannensis by Compton (1 922) and subsequently known as T. elongata was incorrectly 
reduced to synonymy under T. vieillardii. Chinnock (1975, 1976) showed that T. 
tannensis s.str. and T. elongata s. str. are both confined to New Zealand and Australia 
and referredthe NewCaledonian materialtothetaxon he described from New Zealand 
as T. sigmatifolia. Brownlie's T. vieillardii also included a third element, which is a 
member of the T. truncatum R.Br, group. This rather distinctive taxon has never been 
formally recognised taxonomically but was illustrated by Bierhorst (1971). It is 
described here as a subspecies of T. oblanceolata Copel., a Pacific species already 
recorded by the author (Braithwaite 1986) from New Caledonia. 

Four species are, therefore, considered here to occur in New Caledonia; T. 
lanceolata, T. sigmatifolia, T. oblanceolata and T. vieillardii. Since there is no recent 
systematic account covering all species present on the island, the cytological and 
anatomical observations will be presented in the context of a brief taxonomic 
treatment. 

MATERIALS AND METHODS 

Plants of T. lanceolata, T. sigmatifolia and T. oblanceolata subsp. linear if olia were 
collected during a brief visit to New Caledonia in November 1971. Material of each 
species was pressed and dried for herbarium specimens and also preserved in 70% 
alcohol for anatomical studies. Synangia were fixed in the field in 1 :3 acetic alcohol 
and despatched by air to the U.K. for storage in deep freeze. 

The material for anatomical studies was embedded in paraffin wax and the 
sections stained in safranin and light green. Meiotic preparations from the synangial 
material were made usingtheacetocarminesquash method. Sporesfor measurement 
were taken from dried specimens and mounted in gum chloral. Voucher specimens are 
deposited at the Herbarium, Royal Botanic Gardens, Kew (K). 



KEY TO SPECIES AND SUBSPECIES 

1 Sterile leaves distichously arranged, ovate to ovate-oblong with acute apices; sporophylls 
restricted to base or lower half of leafy shoot T. lanceolata 

V Sterile leaves spirally arranged, linear or narrowly oblong to oblanceolate with obtuse 
rounded or truncate apices; sporophylls usually in the middle or upper two thirds of leafy 
shoot 2. 

2 Leaves sigmoid with distal ends incurved 3. 

2' Leaves straight or falcately recurved 4. 

3 Plants epiphytic on tree'fern trunks; leaves narrowly oblong (l/b ratio <5) usually broadest 
below the middle, apices truncate with mucro in the middle T. sigmatifolia 

Plants terrestrial; leaves linear or very narrowly oblong (l/bratio>5) with parallel sides 
or broadest above the middle, apices sharply incurved and rounded with mucro to one 
side almost at right angles to leaf T. oblanceolata subsp. linearifolia 

4 Leaves narrowly oblong to oblanceolate, more or less straight 

T. oblanceolata subsp. oblanceolata 

4' Leaves linear or narrowly oblong but never oblanceolate, regularly falcately recurved 

T. vieillardii 



TMESIPTERIS IN NEW CALEDONIA 201 



OBSERVATIONS ON THE SPECIES 

Tmesipteris lanceolata Dang., Le Botaniste II: 214 (1890-91); Compton, J. Linn Soc. 
Bot. 45: 461 (1922); Brownlie, Flore de la Nouvelle-Cal^donie et D^pendances 3, 
Pte>idophytes: 12, pl.1,f. 1 & 2(1969); Bierhorst, Morphology of Vascular Plants : 154, 
f. 12-1D & E (1971); Chinnock, N.Z. J. Bot. 13 : 759(1975). 

T. tannensis (Spr.) Bernh. var. lanceolata (Dang.) Sahni, Trans. Roy. Soc. Lond. 

ser. B, 213 : 145, pi. 5, f. 9 & 10(1925). 

T. tannensis (Spr.) Bernh. var. typica Sahni, Trans. Roy. Soc. Lond. ser. B, 213 : 

145, pi. 5, f. 8(1925). 
A well known species recognised by all recent authors and distinguished by rat her stiff 
ovate to ovate-oblong leaves with acute apices, sporophylls confined to the base or 
lower half of the leafy shoot and by the distichous arrangement of the leaves beyond 
the sporophylls. It is a common epiphyte usually on the trunks of Cyathea and 
Dicksonia species but is also found on other trunk forming fern genera (see below) 
which presumably provide a similar substrate. 

Stem anatomy: previously investigated by Dangeard (I.e.) and Sahni (I.e.) who both 
reported a stele with a sclerenchymatous pith. The results of the present investigation 
confirm the earlier studies. A section of the aerial shoot at the top of the transition 
region (Fig. 2a) shows the stele made up of three or four groups of tracheids 
surrounding a small and rather ill defined pith composed ofthickwalled lignified cells. 
The pith cells, though not always clearly distinguishable from the tracheids and 
phloem elements in transverse sections, can be readily recognised in longitudinal 
sections by the oblique slit like pits in their side walls. 
Chromosome number: n = 1 04 (Fig. 1a,b). 
Spore size: (51-) 56-58 (-63)pm long, (18-) 21-22 (-25)/um broad. 

Specimens examined: Mt. Koghi, epiphyte on Cyathea trunks, 800m, 1 1 Nov. 1 971 , Braithwaite 
537 (BM, K); Mt. Koghi, epiphyte on trunks of Orthiopteris firma (Kuhn) Brownl., 750m, 1 1 Nov. 
1971, Braithwaite 540 (BM, K). 

Tmesipteris sigmatifolia Chinnock, N.Z. J. Bot. 13: 764 (1975). 

T. tannensis (Spr.) Bernh. auct. Compton, J. Linn. Soc. Bot. 45: 462 (1922). 
T. tannensis (Spr.) Bernh. var. elongata Dang. auct. Sahni, Trans. Roy. Soc. Lond. 
ser. B, 213: 145, pi. 5, f. 5 & 6 (1925). 

T. vieillardii Dang. auct. pro parte Brownlie, Flore de la Nouvelle-Cal^donie et 
D^pendances 3, Pte>idophytes: 12 (1969). 

An epiphytic species up to 20cm long with sigmoid narrowly oblong leaves which are 
incurved at the distal end. It appears to be confined exclusively to tree ferns. 

Stem anatomy: Sahni (1 925) studied the anatomy of the aerial shoot and found it 
essentially similar to that of T. lanceolata. A transverse section from the top of the 
transition region is shown in Fig. 2b. The stele here is made up of a ring of five or six 
tracheid bundles and their associated phloem surrounding a clearly defined pith. The 
pith cells are thick walled and lignified and in longitudinal sections show oblique slit 
like pits characteristic of fibres. The pith is generally. larger and more clearly defined 
than in T. lanceolata. 

Chromosome number: n = 1 04 (Fig. 1c, d). 

Spore size: No mature spores were available for measurement. 

Specimen examined: Montague des Sources, epiphyte on trunks of Dicksonia baudoumi 
Fourn., 900m, 10 Nov. 1971, Braithwaite 529 (BM, K). 



202 FERN GAZETTE: VOLUME 1 3 PART 4 (1 988) 






4b 









FIGURE 1 . Permanent acetocarmine preparations of meiosis. X 750. a) T. lanceolata. Braithwaite 
537. b) Explanatory diagram showing 104 bivalents. c) T. sigmatifolia, Braithwaite 529. 
d) Explanatory diagram showing 104 bivalents. 



Tmesipteris oblanceolata Copel., Philip. J. Sci. 60: 99 (1936); Braith., Brit. Fern Gaz. 

1 0: 296 (1 973); Braith., Fern Gaz. 1 3: 94 (1 986). 

subsp. oblanceolata. 

Plants terrestrial or epiphytic on angiospermous trees, 15- 38cm long. Leaves 10- 16mm long, 
2. 5-3. 5mm broad, narrowly oblong or rectangular with a truncate sometimes bilobedmucronate 
apex to narrowly obovate with rounded mucronate apex. Chromosome number n - 208. 

Distribution: Solomon Islands, New Caledonia, Vanuatu, Fiji, Samoa. 



TMESIPTERIS IN NEW CALEDONIA 



203 




m * -J, 









^^SrN 






^ 






FIGURE 2. Transverse sections of aerial shoots of Tmesipteris species from New Caledonia. 
X 100. a) T. lanceolata, Braithwaite 540. b) T. sigmatifolia. Braithwaite 529. c) 7". oblanceolata 
subsp. linearifolia subsp. nov., Braithwaite 519. 





FIGURE 3. T. oblanceolata subsp. linearifolia subsp. nov., Braithwaite 519. X 1/2. Specimens 
preserved in alcohol. 



204 FERN GAZETTE: VOLUME 1 3 PART 4 (1 988) 



subsp. linearifolia A. Braith. subsp. nov. Fig. 3. 

T. vieillardii Dang. auct. pro parte, Brownlie, Flore de la Nouvelle-Cale'donie et 

D^pendances, 3 Pte>idophytes: 12 (1969). 

T. sp. Bierhorst, Morphology of Vascular Plants, 154, f. 12-1B & C (1971). 

Plantae terrestes, erectae, 8- 20cm longae. Folia 12- 20mm longa, 1.5- 2.5mm lata, 

lineares, minusve sigmoidea, interdum curvata vel tortilia irregulariter apicibus incurvatis 

rotundatis mucronatis; mucro setaceus, apicem surculi versum ad angulum 90. 

Chromosomatum numerus gametophyticae c.208. 

Holotype: Plaine des Lacs, near tributary of Kuelini River, locally frequent growing out of 

moss on boulders and on forest floor, c. 240m, 9 Nov. 1971, Braithwaite 519 (K). 

Plants terrestrial with rhizome creeping among moss on forest floor and on boulders. 
Aerial shoots erect, (8-) 11-18 (-20)cm long maturing in a single growing season and 
ending in a small leaf like appendage or if fertile in a small sporophyll; transition region (4-) 
5- 7(-8)cm long; leaves and sporophylls spirally arranged, sometimes curvedor twisted in 
different directions giving aerial shoot a rather irregular appearance and tending to 
decrease in length towards the apex. Leaves 12-20cm long, 1.5- 2. 5cm broad, lineartovery 
narrowly oblong, slightly sigmoid, apex incurved and rounded with mucro to one side 
almost at right angles to the leaf and directed towards the apex of the aerial shoot. 
Sporophylls in upper half to two thirds of the leafy shoot, equal in length to leaves but 
becoming gradually reduced towards the apex. Synangia 3-4mm long, 1.5- 2.0mm high, 
persistent, bilocular with sporangial lobes more or less equal. Spores monolete. concavo- 
convex, (70-) 78 (-87fcum long, (26-) 31 (-36)um broad. 

Stem anatomy: transverse sections from the top of the transition region (Fig. 2c) 
show the stele with a ring of 6-8 tracheid bundles surrounding a clearly defined pith 
composed of thin walled non-lignified essentially parenchymatous cells. This section 
is very similar to that illustrated by Bierhorst (1971, Fig. 12-1 1D)from "about halfway 
up the frond" of his Tmesipteris sp. 

Chromosome number: n = c. 208 (Figs. 4, 5). Only a very small quantity of 
synangial material was available for chromosome preparations and this yielded a few 
analysable, though less than perfect, cells. The interpretation of the cell illustrated 
shows 208 bivalents at meiosis, but there are two groups with overlapping 
chromosomes which are difficult to interpret with any degree of confidence. Complete 
accuracy for the number cannot therefore be claimed; nevertheless the error if any is 
not great and the order of the number is not in any doubt. 

Notes: T. oblanceolata subsp. linearifolia is a generally smaller plant than subsp. 
oblanceolata with narrower linear leaves which are usually more or less sigmoid and 
sometimes curved or twisted in different directions giving the aerial shoots a rather 
irregular appearance. The rounded leaf apices with the mucrotoonesideanddirected 
towards the apex of the aerial shoot are also characteristic. Field observations and 
ecological data on herbarium sheets indicate that it is nearly always terrestrial on the 
forest floor or growing on moss covered boulders. 

This new subspecies belongs to the T. truncatum R.Br, group and may represent a 

consistently terrestrial form of the Australian species. It is, however, considered 

preferable for the time being to place it here under T. oblanceolata for the same 

reasons as those put forward by Braithwaite (1 986) for referring material allied to T. 

truncatum from Vanuatu to the Pacific species. 

Other specimens examined: Auf den Bergen bei Paita, 1250m, 5 Oct. 1902, Schlechter 
1 4940 (K); Prope ortans fl. N'Go, 300m, Oct. 1912, Franc 1 52 (K); Source de la N'Go, 300m, Franc 
468 (BM, K); Rain forest north east of St. Louis Mission, found growing from base of moss covered 
angiosperm tree, also in leaf base armour of low tree ferns and in other nearby places from 
ground litter, 540m, 20 Oct. 1947, Bucholz 1 244 (K); Contrefortouestdu mdMaoya. Audessusde 
la Mine Emma, foret humide, plante terrestre dress6\ 1 350m, 1 1 July 1 965, McKee 1 2963 (K). 

Tmesipteris vieillardii Dang., Le Botaniste 2: 212, t.9 & 10 (1890-91); Compton, J. 
Linn. Soc. Bot. 45:462(1922); Sahni, Trans. Roy. Soc. Lond. ser. B, 213: 143-170, pi. 
5, f. 1 & 2 (1925); Brownlie, Flore de la Nouvelle-Cal^donie et De'pendances, 3 
Pte>idophytes: 12 (1969) pro parte; Bierhorst, Morphology of Vascular Plants, 154, f. 
12-16, 12-17A (1971). 



TMESIPTERIS IN NEW CALEDONIA 



205 




FIGURE 4. Permanent acetocarmine preparation of meiosis in T. oblanceolata subsp. linearifolia 
subsp. nov. X 750. 



At A* t * * f 



****** »• ^^ 






XC^ 



FIGURE 5. Explanatory diagram for Figure 4 showing c. 208 bivalents. 



206 FERN GAZETTE: VOLUME 1 3 PART 4 (1 988) 



This very distinctive terrestrial species, which has been recognised by all authors, 
is easily distinguished from all other Tmesipteris species in New Caldeonia by its 
robust habit and numerous rather short stiff regularly falcately recurved leaves. 

Stem anatomy: Dangeard (1890-91) and Sahni (1925) have both described and 
illustrated the anatomy in some detail. A typical transverse section of the aerial shoot 
shows a stele made up of a ring of tracheid bundles surrounding a well defined pith 
consisting of non lignified collenchymatous or parenchymatous cells. 

Chromosome number: No material was available duringthepresent investigation 
for the determination of its chromosome number. It is, however perhaps significant 
that both Sahni ( 1925) and Bierhorst(1 971) reported abnormal spore production in the 
form of mis-shapen or abortive spores, a phenomenon confirmed by the author's own 
observations on herbarium specimens. The abnormal spore production could be a 
metabolic effect caused by temporarily adverse environmental conditions and may not 
be particularly significant. On the other hand it does raise the possibility of some 
permanent meiotic disturbance which could be the consequence of hybridisation. 
Cytological observations would therefore be of particular interest and may ultimately 
be necessary for a proper understanding of this species. 

DISCUSSION 
The chromosome numbers recorded here for three New Caledonian species fall into 
the same cytological pattern as those previously reported by Barber (1957) from 
Australia (and New Zealand) and Braithwaite (1 986) from Vanuatu i.e. of a polyploid 
series based on x = 52 (Lovis 1 977). Thus T. lanceolata and T. sigmatifolia with n = 1 04 
are tetraploids and T oblanceolata subsp. linearifolia with n = c. 208 is an octoploid. 

A total of eleven species of Tmesipteris have now been investigated cytologically 
(Table 1) and of these, 7 are tetraploids and 4 are octoploids, but as yet no diploids have 
been found. Although a consistent cytological pattern for the genus appears to be 
emerging, it should be noted that the sampling of some species is inevitably limited by 
their scarcity and that others have not been sampled at all. More data are desirable 
particularly from New Zealand, New Guinea and the more remote Pacific islands. 

The stelar anatomy of the aerial shoots of the species from New Caledonia is 
basically similar in so far as the stele is made up of a variable number of groups of 
tracheids surrounding a usually clearly defined pith. There are, however, marked 
differences in the nature of the pith cells. The pith of the two tetraploid species, 7". 
lanceolata and T. sigmatifolia, is made up of thick walled lignified cells resembling 
sclerenchyma while that of the octoploid T. oblanceolata subsp. linearifolia is 
composed of non- lignified thinner walled parenchymatous eel Is. The last non- lignified 
type has also been reported in T. vieillardii\Nh\ch has not been studied in detail during 
the present investigation. 

Variation in the type of pith cells in aerial shoots of Tmesipteris has been reported 
by a number of previous investigators. Dangeard (1 890-91) dividedthegenus into two 
sections "selons qu'ils ont des fibres medullaires ou sont depourvus ...", i.e. intothose 
species possessing a sclerenchymatous pith (T. lanceolatum, T truncatum) and those 
species with a parenchymatous or collenchymatous pith (T. vieillardii, T. tannensis) or 
no pith at all (T. elongatum). In the latter case the tracheid bundles were reported to be 
more or less united in the centre of the stele. Sahni (1925) and Braithwaite (1973, 
1986) recognised two basic types of pith composed of either lignified 
sclerenchymatous cells or non-lignified parenchymatous/collenchymatous cells. The 
different pith types reported in the literature are summarised in Table 1 together with 
the ecology of the species. 

The ecological observations presented here on the cytologically authenticated 
material from New Caledonia generally confirm earlier observations in the literature 



TMESIPTERIS IN NEW CALEDONIA 207 



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on these species. Thus 7". lanceolata and 7". sigmatifolia are confined exclusively to tree 
fern trunks whereas 7". oblanceolata subsp. linearifolia, like 7". vieillardii, is essentially 
terrestrial. Elsewhere the majority of Tmesipteris species are recorded as tree fern 
epiphytes but there are a few taxa which are terrestrial, like T. oblanceolata subsp. 
linearifolia and T. vieillardii, or epiphytic on angiosperms (Table 1). The most notable of 



208 FERN GAZETTE: VOLUME 1 3 PART 4 (1 988) 



these are T. tannensis s. str. from New Zealand (Chinnock 1975) and T. oblanceolata 
subsp. oblanceolata from the Solomon Islands (Braithwaite 1973) and Vanuatu 
(Braithwaite 1 986). These species may occasionally be found on tree fern trunks but 
they appear to occur predominantly on non-tree fern substrates. 

The data summarised in Table 1 does not suggest any correlation between the 
chromosome number and either the anatomy or the ecology of the species. The 
tetraploid species have either a sclerenchymatous pith or non-lignified 
parenchymatous/collenchymatous pith, with the former type predominant, and are 
found on tree fern and non-tree fern substrates. Both anatomical and ecological types 
are also found among the octoploid species. There is perhaps a closer, though not 
complete, correlation between pith type and ecology of the species. All those species 
listed in Table 1 growing on tree fern trunks, apart from T. billardieri, have a 
sclerenchymatous pith and those on non-tree fern substrates, whether as an 
angiosperm epiphyte or terrestrial fern, have a non-lignified parenchymatous/collen- 
chymatous pith. The significance of this observation is at present not clear. It may 
merely be an effect of inadequate sampling. On the other hand it could be related 
somehow to the size of the plants or to the different functional requirements of aerial 
shoots in the terrestrial or angiosperm epiphytic environments as opposed to the 
rather specialised environment of tree fern trunks. It is not possible to reach a firm 
conclusion from the present incomplete data. Further studies of species not included 
in Table 1 are needed as well as confirmation of some of the older observations on 
some of the species listed there. 

ACKNOWLEDGEMENTS 
I am grateful to MonsieursSchmid and Vieillon of Ormston, Noumea, fortheir valuable 
assistance during my short visit to New Caledonia and to Mr. B.V. Case, Botany 
Department, University of Nottingham, for technical help and guidance during 
preparation of the illustrations. 

REFERENCES 

BARBER, H.N. 1957. Polyploidy in the Psilotales. Proc. Linn. Soc. N.S.W., 82: 201-208. 
BIERHORST, D.W. 1971, Morphology of Vascular Plants. MacMillan, New York. 
BONAPARTE, R. 1921. Notes Pte>idologiques, Fasc. 3. 

BRAITHWAITE, A.F. 1973. Tmesipteris in the Solomon Islands. Brit. Fern Gaz. 10: 293-303. 
BRAITHWAITE, A.F. 1986. Tmesipteris in Vanuatu (New Hebrides). Fern Gaz. 13: 87-96. 
BROWN, R. 1 806. Prodromus Florae Novae Hollandiae etlnsu/ae vanDiemen. Johnson, London. 
BROWNLIE, G. 1 969. Flore de la Nouvelle-Cale'donie et DSpendances 3: Ptdridophytes. 307pp. 

Museum National d'Histoire Naturelle, Paris. 
CHINNOCK, R.J. 1975. The New Zealand species of Tmesipteris (Psilotaceae). N.Z. J. Bot. 13: 

743-768. 
CHINNOCK, R.J. 1976. The identification, typification and origin of Tmesipteris tannensis 

(Psilotaceae). Taxon 25: 115-121. 
DANGEARD, PA. 1890-91. Memoire sur la morphologie & I'anatomie des Tmesipteris. Le 

Botaniste, Series II: 163-222. 
DIELS, L. 1 906. Filicales, in Schlechter, R. Beitrage zur Kenntnis der Flora von Neu-Kaledonien. 

Bot. Jahrb. fur Syst. 39: 2- 1 3. 
ENDLICHER, S. 1833. Prodromus Florae Norfo/kiae. Wien. 
JARMAN, S.J., KANTVILAS, G. & BROWN, M.J. 1 986. The ecology of pteridophytes in Tasmanian 

Cool Temperate Rainforest. Fern Gaz. 13: 77-86. 
JEANPERT, H.E. 191 1. Fougeres recueilles en Nouvelle-Cal^donie par M et Mme Le Rat et aux 

Nouvelle Hebrides par Mme Le Rat. Bull Mus. d'Hist. Nat. Paris 17: 571-579. 
JEANPERT, HE. 1912. Fougeres de Nouvelle-Cal^donie, r£colt£es par M Cribs. Bull. Mus. d'Hist. 

Nat. Paris 18: 102-107. 
LOVIS, J.D. 1977. Evolutionary patterns and processes in ferns. Adv. Bot. Res. 4: 229-419. 
METTENIUS, G. 1861. Filices Novae-Caledoniae. Ann. Sci. Nat. Ser. IV 15: 55-88. 
SAHNI, B. 1925. On Tmesipteris vieillardii Dangeard, an erect terrestrial species from New 

Caledonia. Phil. Trans. Roy. Soc. B, 213: 143-170. 
SYKES, M.G. 1908. The anatomy and morphology of Tmesipteris. Ann. Bot. 22: 63-89. 
WAKEFIELD, N.A. 1943. Two new species of Tmesipteris. Vict. Nat. 60: 142-143. 



FERN GAZ. 13(4)1988 209 



SHOOT TEMPERATURE MEASUREMENTS OF MONTANE 

CYATHEA (CYATHEACEAE: PTERIDOPHYTA) SPECIES 

IN PAPUA NEW GUINEA 

M.J. EARNSHAW 
Department of Cell and Structural Biology, Williamson Building, 
University of Manchester, Manchester M13 9PL, Great Britain 

T.C. GUNN 

Biology Department, Oakham School, Oakham, Rutland, 

Leicestershire LE15 6DT, Great Britain 

J.R. CROFT 

Division of Botany, Department of Forests, P.O. Box 314, Lae, 

Papua New Guinea 

ABSTRACT 

Scales occur on the crozier and dorsal surface of the shoot apex in Cyathea spp. 
and dead frond bases are found on the lateral surfaces of the apex. Diurnal 
temperature measurements were made on plants growing at approx. 3500m. a. s. I. 
in order to determine whether these external structures are effective in protecting 
active tissues from low nocturnal temperatures in the tropicalpine environment. 

At dawn air temperatures of 0-2°C, the croziers and pinnae of C. gleichenioides 
and C. atrox var. atrox were generally less than ambient with sub-zero 
temperatures recorded on occasion. Internal shoot apex temperatures, however, 
were consistently found to be 2.0-3. 5°C above air temperature suggesting that 
the apical meristem is protected by surrounding structures from the potentially 
deleterious effect of low nocturnal temperatures. 

Subsequent daytime temperature recordings revealed that the shoot apex and, 
to a lesser extent, the crozier tissue were slow to equilibrate to increasing air 
temperature. Croziers were substantially shaded by mature fronds but on receipt 
of direct irradiation, at around midday, showed an increase in temperature of approx. 
5°C above ambient. Such a temperature increase will enhance the rate of crozier 
growth and development within this low temperature environment. 

INTRODUCTION 
The alpine regions of tropical high mountains are exposed to a high amplitude of 
diurnal temperature change but with little variation from season to season. The "giant 
rosette" plants represent an adaptation to this environment with representative 
examples in the genera Espeletia in the Andes and Dendrosenecio, Lobelia and Senecio 
occurring in the East African mountains. In these species the apical bud is surrounded 
by young developing leaves which show nyctinastic leaf movement, apparently serving 
to protect the shoot apex from low nocturnal temperatures which are frequently sub- 
zero (Smith 1974; Larcher 1975; Hedberg & Hedberg 1979; Beck et al. 1982; Rada 
et al. 1985). In addition, the layer of dead leaves which remain attached to the stem 
in Espeletia provides thermal insulation thus preventing freezing temperatures from 
reaching active tissues (Smith 1979; Goldstein & Meinzer 1983; Rada et al. 1985). 
The occurrence of this life-form on two widely separated mountain ranges is frequently 
considered to be an example of convergent evolution. 

"Giant rosette" plants, however, are not found in the Papuasian mountains but 
Cyathea spp. occur commonly in exposed sub-alpine grassland to an altitude of approx. 
3800m. a. s.l. where sub-zero nocturnal temperatures can occur (Wade & McVean 
1969; Hnatiuk et al. 1976; Hope 1980). The lateral surfaces of the shoot apex are 
composed of dead frond bases and dense scales clothe both the flat dorsal surface 
of the apex and the young developing fronds. The measurements reported in this 
paper were planned to determine the effectiveness of these external structures in 
insulating active tissues from low nocturnal temperatures. 



210 



FERN GAZETTE: VOLUME 1 3 PART 4 (1 988) 



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FIGURE 1. Pindaunde Valley, Mt Wilhelm. The inset of Papua New Guinea shows the major 
mounta.n ranges (after Hnat.uk et al. 1 976 with altitudes from Humphreys 1 984). 



CYATHEA IN PAPUA NEW GUINEA 



211 



MATERIALS AND METHODS 
The study was conducted on Mt. Wilhelm, Papua New Guinea near the Pindaunde 
Research Station (5°47'S, 145°03'E; 3580m. as. I.) as depicted in Fig. 1 . The Pindaunde 
Valley is a hanging, U-shaped valley located on the south eastern flank of Mt Wilhelm. 
Climatic records for Pindaunde Research Station (Hnatiuk et al. 1976) show mean 
annual minimum and maximum temperatures of 4.0°C and 11.6°C respectively at 
screen height (1.5m above ground) with infrequent sub-zero temperatures recorded. 
However, the ground surface in open areas has a more rigorous temperature regime 
with mean annual minimum and maximum temperatures of about 1.1 °C and 19 7°C 
and a minimum recorded extreme of -9.1°C. Freezing air temperatures are also 
common at canopy level (0.5-1 0m above ground) in exposed situations. 

Taxonomy of the tree fern spp. involved in the present study is that of Holttum 
(1963) and Holttum & Edwards (1983). The vegetation of the Pindaunde Valley (Fig. 
1 ) consists of sub-alpine tussock grassland, often with extensive tree fern shrublands, 
and sub-alpine rainforest which is confined to the valley sides (Wade & McVean 
1969). The lower part of the Valley at Kombuglomambuno contains the montane 
tree fern alliance as described by Hope (1980) and is dominated by Cyathea atrox 
C.Chr. which also occurs commonly in the forest. The sub-alpine tree fern alliance 
(Hope 1 980) occurs at approx. 3400-3800m.a.s.l. and is dominated by C. gleichenioides 
C.Chr. and C. muelleri Bak. which are generally not found in the forest. 

Detailed diurnal time course measurements were made of shoot temperatures 
in single plants of C. gleichenioides and C. atrox var. atrox (see Fig. 1 and Table 
1) generally at 30 minute intervals. Irradiance readings were taken with a miniature 
tube solarimeter (Delta-T, Cambridge, UK) previously calibrated with a Kipp and Zonen 
"Solarimeter". Total irradiance was determined in an unobstructed position in the 
open and irradiance reaching the croziers and dorsal surface of the shoot apex 
measured by placing the solarimeter at a height of approx. 20cm above the apex. 
Crozier and pinna surface temperatures were recorded using copper-constantan 
thermocouples (Comark, Rustington, Littlehampton, West Sussex, UK) linked to Wescor 
HR33T microvoltmeters (Wescor Instruments, Logan, Utah, USA). Crozier tissue 
temperature was recorded using a fixed thermocouple inserted into the tissue in the 
centre of the terminal coil at least 24 hours prior to taking readings. The croziers 
utilised were 9cm in height in the case of C. gleichenioides (Fig. 2) and 2cm in height 
for C. atrox var. atrox (Table 2). Internal shoot apex temperatures were measured 
with thermistors (Edale thermistor Model K, Edale Instruments (Cambridge) Ltd., UK). 
Thermistors were located at least 24 hours prior to reading in the centre of the dorsal 
surface of the apex and/or laterally approx. 1 0cm below the dorsal surface. Thermistor 
depth was 5cm for C. gleichenioides (Fig. 2) and 9cm for C. atrox var. atrox 
(Table 2). 

TABLE 1. Location and characteristics of the Cyathea individuals utilised in this study (see 
also Fig. 1). 





C. GLEICHENIOIDES 


C. ATROX var. ATROX 


Altitude, m.a.s.l. 


3470 


3500 


Location 


Tussock grassland 


Rainforest edge 


Aspect 


Fully exposed 


Fully exposed 


Shoot height, cm 


102 


410 


Shoot apex diameter, cm 


26 


29 


Mature fronds per plant 


21 


12 


Mature frond length, cm 


94 


174 



212 



FERN GAZETTE: VOLUME 1 3 PART 4 (1 988) 



1500 



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20 

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14 AUGUST, 1984 



TIME IN HOURS 



20 AUGUST, 1984 



FIGURE 2. Shoot temperature recordings of the same individual of Cyathea gleichenioides on 
a cloudy cool day (A) and on a clear warm day (B). Solid lines on the graphs of shoot temperature 
depict air temperature. For the sake of clarity, data points for pinna temperature have been 
omitted on the cool day (A) except for the period of maximum irradiance. Minimum ground 
surface temperatures recorded at 6.00 hours were -3.0°C (A) and -1 .0°C (B). 



CYATHEA IN PAPUA NEW GUINEA 



213 



TABLE 2. Sample shoot temperature recordings of Cyathea atrox var. arrox on a clear warm 
day (22 August, 1 984). The data are taken from a detailed time course as in Fig. 2. The minimum 
ground surface temperature recorded was -1 .0°C at 6.30 hours. 



TIME 


IRRADIANCE, Wm 2 


TEMPERATURE RECORDINGS, °C 


Hours 


Total 


Shoot apex 


Air 


Shoot apex 


Pinna 


Crozier 
surface 


Crozier 
tissue 


6.00 
10.00 
14.00 




720 

1360 



380 

945 


0.1 
11.7 
17.0 


3.5 

7.0 

16.5 


0.1 
12.5 
17.0 


-0.3 
13.0 
22.0 


-0.4 

8.9 

22.2 



RESULTS 



Figure 2 depicts temperature recordings of various shoot tissues of a single individual 
of C. gleichenioides on a cloudy cool day and on a clear warm day. 

Irradiance measurements on the cool day (Fig. 2A) showed shading by mature 
fronds of the dorsal surface of the shoot apex and attendant croziers. The internal 
temperature of the shoot apex at dawn was greater than air temperature by 2.5- 
3.5°C. Thereafter, the shoot apex slowly reached a maximum temperature in the 
afternoon, which was rather higher underneath the dorsal surface, but generally 
remained below ambient temperature. During the evening, the decline in shoot apex 
temperature was much less rapid than the fall in air temperature (see also Fig. 2B). 
Surface temperatures of the crozier were equal to or less than ambient throughout 
the day but the post-dawn increase in crozier tissue temperature lagged behind the 
increase in air temperature. Unexpectedly, the brief period of higher irradiance produced 
an increase in crozier tissue temperature of approx. 4°C above crozier or pinna surface 
temperature. 

A further set of measurements was taken on a warm day (Fig. 2B) in order to 
investigate further the midday increase in crozier tissue temperature. Figure 2B shows 
that the crown and young croziers were substantially shaded by the surrounding 
mature fronds except for a period around midday (approx. 11-14 hours). Nocturnal 
and dawn temperatures of the pinna and crozier were generally less than air 
temperature in contrast to internal shoot apex temperatures which were well above 
ambient. Interestingly, the maximum shoot apex temperature was little higher on 
the warm day (Fig. 2B) than that occurring on the cool day (Fig. 2A). Morning 
temperatures of the crozier and pinna were mostly above air temperature on the 
warm day (Fig. 2B). The midday period of more or less unobstructed irradiance at 
the shoot apex increased crozier temperatures substantially above that of the air or 
pinna surface with the crozier surface temperature being higher than the tissue. 

Similar determinations were also carried out with C. atrox var. atrox on a clear 
warm day (Table 2) and confirm the major findings for C. gleichenioides on a similar 
day (Fig. 2B). The internal shoot apex of C. atrox var. atrox, but not the crozier, was 
protected against low nocturnal temperatures and the post-dawn increase in shoot 
apex and crozier tissue temperature showed a lag with respect to ambient. However, 
the shoot apex of C. atrox var. atrox attained a higher maximum temperature (Table 
2) than C. gleichenioides (Fig. 2B) probably due to a reduced density of scales on 
the dorsal surface producing a less well-insulated structure. A similar shading effect 
of the shoot apex of C. atrox var. atrox occurred as in C. gleichenioides (Fig. 2B) 
but the midday period of high irradiance caused an increase in crozier temperature 
of approx. 5°C above air or pinna temperature (Table 2). 



214 FERN GAZETTE: VOLUME 1 3 PART 4 (1 988) 



DISCUSSION 

The most unexpected finding in this work was the increase in crozier temperature 
of approx. 5°C above ambient in response to direct irradiation (Fig. 2 and Table 2), 
a feature previously observed in other bulky plant organs. Daytime temperatures of 
leaf and stem succulents can rise up to 20°C above air temperature giving rise to 
tissue temperatures as high as 60°C (Patten & Smith 1975; Nobel 1978; Larcher 
1980; Kappen 1981). As the degree of succulence increases there is a decrease 
in the rate of heating but a greater retention of energy (Mellor et al. 1 964). An increase 
in daytime temperature also occurs in the apical region of Espeletia which has been 
ascribed to the parabolic configuration of the leaf rosette thus concentrating incoming 
radiation to the centre (Smith 1974; Larcher 1975; Hedberg & Hedberg 1979). Recent 
work on Espeletia schultzii\Ne66. has demonstrated an increase in the vertical 
orientation of the central rosette leaves at higher altitudes together with a more 
pronounced curvature of the leaf bases towards the main axis (Meinzer et al. 1985). 
It was suggested that this morphological change may enhance the capture and retention 
of incident radiation by the apical bud leading to more favourable temperatures for 
apical growth and leaf expansion. 

Temperature increases in the above bulky plant organs arise as a result of a 
large heat capacity relative to a small surface area available for heat dissipation. 
In comparison to many succulents, the relatively high surface area: volume ratio of 
the Cyathea crozier produced a relatively fast heating rate but only limited energy 
retention (Fig. 2). In ferns, crozier uncoiling involves elongation of the rachis and 
differential cell elongation on the abaxial and adaxial surfaces and is mediated by 
polar transport of auxin produced in the pinnae (Briggs & Steeves 1 958, 1 959; Steeves 
& Briggs 1960; Voeller 1966). It seems likely that enhanced daytime temperatures 
of the crozier of Cyathea spp. will enhance the rate of crozier development in plants 
growing in this low temperature environment. 

Nocturnal temperatures of both the crozier and pinna of Cyathea spp. tended 
to be lower than ambient and on occasion were sub-zero (Fig. 2 and Table 2). These 
reduced tissue temperatures were presumably due to energy exchange with the 
surrounding air via long-wave infra-red emission which is a common feature of the 
tropicalpine environment (Beck et al. 1 982; Rada et al. 1 985). The scales of the crozier, 
therefore, appear to play no part in protecting the tissue from low nocturnal 
temperatures. Similarly, the pubescent layer on the lower leaf surfaces surrounding 
the night-bud in some "giant rosette" plants does not decrease infra-red emission 
(Beck et al. 1982; Rada et al. 1985). 

By contrast, the microenvironment of the large shoot apex of mature sporophytes 
of Cyathea spp. was relatively constant in temperature, particularly in the case of 
C. gleichenioides (Fig. 2), and protected from low nocturnal temperatures (Fig. 2 and 
Table 2). Stem temperatures of Espeletia spp. also remain above air temperature 
during the night with marked insulation being provided by the dead leaves (Smith 
1979; Goldstein & Meinzer 1983; Rada et al. 1985). Experimental removal of the 
layer of dead leaves increases plant mortality which appears to be due to the effect 
of sub-zero temperatures on water availability (Smith 1979; Goldstein & Meinzer 
1983). Although sub-zero nocturnal air temperatures at heights >1m above ground 
level were not recorded in the Pindaunde Valley in the present study (Fig. 2 and 
Table 2), freezing air temperature below this height are common (Hnatiuk et al. 1 976). 
Young Cyathea sporophytes clearly have to contend with a more rigorous temperature 
regime than mature individuals implying that these species must possess some degree 
of resistance to freezing stress. Andean "giant rosette" plants avoid freezing by means 



CYATHEA IN PAPUA NEW GUINEA 215 



of supercooling and do not tolerate ice formation within the tissues (Larcher 1981; 
Rada et al. 1985). On the other hand, the Afroalpine species are able to tolerate 
freezing to the extent that full photosynthetic capacity is regained immediately after 
thawing (Beck et al. 1982, 1984; Schulze et al. 1985). A comparable investigation 
into freezing resistance of Papuasian Cyathea spp. clearly needs carrying out. 



ACKNOWLEDGEMENTS 
This work was carried out as part of the Oakham School Expedition to Papua New 
Guinea, 1984 and our thanks are due to all those who assisted with everyday duties 
and obtaining the data presented. Permission to use Pindaunde Research Station 
was kindly granted by the National Parks Service, Office of Environment and 
Conservation, Port Moresby and the Assistant Director, Division of Botany (Lae) was 
generous in making other facilities available. The staff of the Department of Biology, 
University of Papua New Guinea, Port Moresby provided extensive support and the 
Percy Sladen Memorial Fund financial assistance. We would also like to thank 
Mr D. Tarrant for technical assistance; June Underwood for typing the manuscript; 
Dr D.T. Clarkson, Haverhill Generators (Kettering, UK), Phillip Harris Ltd (Shenstone, 
Staffs, UK), Phillips (UK) Ltd, Dr D. Ratcliffe, Dr J. Roberts and Wescor Instruments 
(Logan, Utah, USA) for the loan or donation of equipment; and Professor E.G. Cutter, 
Dr E. Sheffield and Dr R.A. White for useful advice and discussion. 



REFERENCES 

BECK, E., SENSER, M., SCHEIBE, R., STEIGER, H.-M. & PONGRATZ, P. 1982. Frost avoidance 

and freezing tolerance in Afroalpine "giant rosette" plants. Plant, Cell Environ. 5: 212- 

222. 
BECK, E., SCHULZE, E.-D., SENSER, M. and SCHEIBE, R. 1984. Equilibrium freezing of leaf 

water and extracellular ice formation in Afroalpine "giant rosette" plants. Planta 162: 

276-282. 
BRIGGS, W.R. & STEEVES, T.A. 1958. Morphogenetic studies on Osmunda cinnamomea - the 

expansion and maturation of vegetative fronds. Phytomorphology 8: 234-248. 
BRIGGS, W.R. & STEEVES, T.A. 1959. Morphogenetic studies on Osmunda cinnamomea L. 

-the mechanism of crozier uncoiling. Phytomorphology 9: 134-147. 
GOLDSTEIN, G. & MEINZER, F. 1983. Influence of insulating dead leaves and low temperatures 

on water balance in an Andean giant rosette plant. Plant, Cell Environ. 6: 649-565. 
HEDBERG, I. & HEDBERG, O. 1979. Tropical-alpine life-forms of vascular plants. Oikos 33: 

297-307. 
HNATIUK, R.J., SMITH, J.M.B. & McVEAN, D.N. 1976. Mt Wilhelm Studies 2. The climate of 

Mt Wilhelm. ANU, BG/4, Canberra. 
HOLTTUM, R.E. 1963. Cyatheaceae. In: Flora Malesiana, Series II, Pteridophyta, Vol. 1(2): pp. 65- 

176. 
HOLTTUM, R.E. & EDWARDS, P.J. 1983. The tree ferns of Mount Roraima and neighbouring 

areas of the Guayana Highlands with comments on the family Cyatheaceae. Kew Bull. 

38. 155-188. 
HOPE, G.S. 1980. New Guinea mountain vegetation communities. In: van Royen, P. The alpine 

flora of New Guinea, Vol. 1. J. Cramer, Vaduz, pp. 1 53-222. 
HUMPHREYS, G.S. 1984. The environment and soils of Chimbu Province, Papua New Guinea 

with particular reference to soil erosion. Research Bulletin 35. DPI, Port Moresby. 
KAPPEN, L. 1981. Ecological significance of resistance to high temperature. In: Lange, O.L., 

Nobel, P.S., Osmond, C.B. & Ziegler, H. (eds). Encycl. Plant Physiol. (NS), 12A, Springer, 

Berlin, Heidelberg, New York, pp. 438-474. 
LARCHER, W. 1975. Pflanzenokologische Beobachtungen in der Paramostufe der 

venezolanischen Anden. Anzeig. Math. - naturw. Kl. Osterr. Akad. Wiss. 1 1: 1-20. 
LARCHER, W. 1980. Physiological plant ecology. Springer, Berlin, Heidelberg, New York. 303pp. 
LARCHER, W. 1981. Physiological basis of evolutionary trends in low temperature resistance 

of vascular plants. Plant Syst. Evol. 137: 145-180. 
MEINZER, F.C., GOLDSTEIN, G.H. & RUNDEL, P.W. 1 985. Morphological changes along an altitude 

gradient and their consequences for an Andean giant rosette plant. Oecologia (Berlin) 

65: 278-283. 



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MELLOR, R.S., SALISBURY, F.B. & RASCHKE, K. 1964. Leaf temperatures in controlled 

environments. Planta 61: 56-72. 
NOBEL, P.S. 1 978. Surface temperatures of cacti - influences of environmental and morphological 

factors. Ecology 59: 986-996. 
PATTEN, D.T. & SMITH, E.M. 1 975. Heat flux and the thermal regime of desert plants. In: Hadley, 

N.F. (ed.). Environmental physiology of desert organisms. Dowden, Hutchinson and Ross, 

Stroudsburg Pa, pp. 1-19. 
RADA, F., GOLDSTEIN, G., AZOCAR, A. & MEINZER, F. 1985. Freezing avoidance in Andean 

giant rosette plants. Plant, Cell Environ. 8, 501-507. 
SCHULZE, E.-D., BECK, E., SCHEIBE, R. & ZIEGLER, P. 1985. Carbon dioxide assimilation and 

stomatal response of Afroalpine giant rosette plants. Oecologia (Berlin) 65: 207-213. 
SMITH, A. P. 1974. Bud temperature in relation to nyctinastic movement in an Andean giant 

rosette plant. Biotropica 6: 263-266. 
SMITH, A.P. 1979. The function of dead leaves in Espeletia schultzii (Compositae), an Andean 

giant rosette plant. Biotropica 1 1: 43-47. 
STEEVES, T.A. & BRIGGS, W.R. 1960. Morphogenetic studies on Osmunda cinnamomea L 

- the auxin relationships of expanding fronds. J. Exp. Bot. 1 1: 45-67. 
VOELLER, B.R. 1966. Crozier uncoiling of ferns. Rockefeller Univ. Rev. 4: 14-19. 
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ANU, BG/1, Canberra. 



FERN GAZ. 1 3(4) 1 988 217 



PTERIDOPHYTES OF ZARATE, A FOREST ON THE WESTERN 
SIDE OF THE PERUVIAN ANDES 



BLANCA LEON and NIELS VALENCIA 

Museo de Historia Natural 

Universidad Nacional de San Marcos 

Casilla 14-0434, Lima 14, Peru. 

ABSTRACT 
Twenty-three species of pteridophytes are reported from an area located on the 
western slopes of the Andes in central Peru, which has been proposed as a nature 
reserve. A taxonomic key is provided. Habitats and microhabitats of the area are 
described. Biogeographical elements are discussed. 

INTRODUCTION 
The vegetation of the western slopes of the Andes in central Peru has been considered 
as semidesert and steppe (Weberbauer 1936, 1945; Tryon 1960, 1971). In 1952, 
H. and M. Koepcke discovered a small forest, called "Zarate", on a very steep lateral 
valley of the Rimac river. Later, the Koepckes found other smaller forests along the 
western slope of the Andes (Koepcke 1954, Koepcke & Koepcke 1958); all of them 
are in steep valleys of difficult access, between 2600 and 3500m elevation. These 
forests were considered to be an extension of the humid temperate zone (Koepcke 
1954), which was first described by Chapman (1926) for Ecuador and Northwestern 
Peru. It is possible that in the past they formed a more or less continuous forest 
belt at that elevation interrupted only by rivers and reaching as far as 16°S (Koepcke 
1961). 

The pteridophyte flora of Zarate forest, as well as the surrounding area, was surveyed 
as part of the research needed to establish the area as a nature reserve. There are 
very few publications about the pteridophytes of western Peru. This study is the first 
done on the pteridophytes of the western slopes of the Andes in central Peru. Tryon 
(1960, 1971) made a brief reference to this area, describing general aspects of the 
environment. Floristic studies done by Sanchez (1965) and Vasquez (1967) refer to 
northern Peru, where climate and topography are different, while that of Leon (1983) 
is limited to the pteridophytes of the central coast below 1000m elevation. 

THE STUDY AREA 
The study area is located in the Department of Lima, between 1 1 °53'1 7"-1 1 °56'1 7"S 
and 76°25'41"-76°31'07"W (Fig. 1). The approximately 10km 2 area lies largely on 
the right side of the Seco river, which is part of the drainage of the Rimac river. 
The valley walls are very steep, with slopes ranging from 40° to vertical cliffs; there 
are several smaller V-shaped ravines. This study includes the Zarate forest, and the 
surrounding area down to 1500m elevation. 

Bedrock in the area is volcanic in origin. Soils in general are immature, acidic, 
and coarsely textured in the lower elevations and in open sites in the forest. The 
percentage of sand in the soil tends to decrease with increasing altitude. In the forest, 
the soils are the most developed, being generally nitrogen rich, and slightly acidic 
or neutral (Franke & Valencia 1984). 

Climatic data are not available for the study area. The general characteristics have 
been deduced from several meteorological stations at different altitudes and more 
or less similar latitudes (Franke & Valencia 1 984). These reveal gradients in temperature 
and rainfall with altitude. Here the rainfall is seasonal; ninety percent of the annual 



218 



FERN GAZETTE: VOLUME 13 PART 4 (1988) 



! 



/ .. < • Z A RATE 




20km 

— I 



Figure 1 . Map of the study area. 

rainfall is concentrated during the months from November to March. In the lower 
parts (1500m) it is probably c. 170mm, and in the forest (2860m) it is c. 360mm. 
During the rainy season (southern summer) heavy mist occurs often in the forest. 
In the lower part of the study area, the mean monthly maximum temperature ranges 
from 19°C in July to 22°C in February; mean monthly minimum temperatures from 
1 1°C in August to 14°C in March. In the forested area, the mean monthly maximum 
temperature ranges from 17°C in February to 18°C in July; mean monthly minimum 
temperature varies from 7°C in July to 9°C in March. 

Most of the study area has been disturbed by human activities. For centuries 
the local population has probably utilized it as a source of forage and fuel. Several 
archeological structures are present in the forest. The lowest elevations, 1500 to 
1800m, are presently used for agriculture. 



PTERIDOPHYTES OF ZARATE 219 



METHODS 
The main hiking trail crosses the altitudinal range of the study area and thus was 
used as a transect about 10km long. Collections were made along the trail, surveying 
roughly 20m to each side of it, from 1977 to 1984, by the authors, I. Franke, and 
A. Cano. These collections are deposited at the Herbario del Museo de Historia Natural 
(USM), Universidad Nacional de San Marcos, Lima. Some specimens were sent for 
identification to J. T. Mickel at the New York Botanical Garden (Elaphoglossum), and 
A. R. Smith at the University of California, Berkeley (Thelypteris). Poorly known species 
with nomenclatural difficulties are treated here by using the most common name 
found in the literature. 

SPECIES LIST 
Thirteen genera, twenty-three species, and three varieties have been found in the 
study area. A taxonomic key is provided in Appendix 1 . 
Adiantum digitatum Hooker 

This species grows from Ecuador to Brazil and northwest Argentina. In Peru it occurs in 
inter-andean valleys and on the western side of the Andes between 400m and 4000m elevation. 
It is found in the forest of the study area in crevices among rocks. 

N.V. & I.F. 23 (2700m, Cheqta, 12 Jun 1977); 1127 (2860m, Gigantdn, 1 May 1981); 1273 
(286m, Gatero, 1 May 1981). 
Adiantum poiretii Wikstr. 

This species occurs in Tropical America and in the Old World tropics. In the study area, 
we found two of the currently recognized varieties for Peru (Tryon 1 964): 
Adiantum poiret /'/'Wikstr. var. poiretii 

This variety occurs from Mexico, Central America and the Antilles to Bolivia and Uruguay. 
In Peru it grows on both sides of the Andes and in inter-andean valleys between 800m and 
3900m elevation. In the study area it is commonly found in humid, shaded parts of the forest. 
N. V. & I. F. 514 (2850m, Mayhuayqui, 13 Jan 1980); 560; and 610 (2880m, Carnacha, 4 
Mar 1980). 
Adiantum poiretii Wikstr. var. sulphureum (Kaulf.) R. Tryon 

This variety grows from Peru to Chile and Argentina, at the same altitude as var. poiretii. 
In the study area it only occurs in humid, shaded sites in the forest. 
N. V. & I. F. 346 (3000m, Carnacha, 25 Mar 1978); 1277 (2860m, Gatero, 1 May 1981). 
Adiantum subvolubile Kuhn 

This species occurs in Ecuador and Peru. In Peru it grows in seasonal coastal vegetation 
("lomas"), on both sides of the Andes and in inter-andean valleys. In the study area it has 
been found among rocks in sites protected by the vegetation. 

N. V. & I. F. 276 (2850m, Gigantdn, 25 Mar 1978); 592 (2870m, Gigantbn, 4 Mar 1980); 1143 
(2000m, Chunaca, 1 May 1980). 
Asplenium aethiopicum (Burm.f.) Becherer (syn. A. praemorsum Sw.) 

This species is widely distributed in the tropics of the Old and New Worlds. In Peru it occurs 
in the departments of Amazonas, Junin, and Cuzco, between 1200m and 3200m elevation. 
In the study area, it grows among rocks and in partially open sites. This is the first report 
of this species on the western side of the central Andes. 
N. V. & I. F. 1 275 (2860m, Gatero, 1 May 1 981 ). 
Asplenium fragile Presl 

This species is distributed from Venezuela to Chile, above 2800 m elevation. In Peru it occurs 
in the departments of Cajamarca, Ancash, Junin, Lima, Huancavelica, Ayacucho, and Puno. 
In the study area it grows among rocks, and in humid, shaded sites in the forest. 
N. V. & I. F. 763 (2870m, Gigantdn, 20 Apr 1 980); 1 526 (31 00m, Pampa Zarate, 23 Apr 1 982). 
Asplenium sessilifolium Desv. 

This species is distributed from Mexico and Central America to Colombia, Peru and Bolivia. 
In Peru it has been found in inter-andean valleys between 2900m and 4000m elevation. In 
the study area, it occurs in humid, shaded sites, sometimes together with A. fragile. 
N. V. & I. F. 145 (3100m, Muralla, 30 Jul 1977); 278 (2850m, Gigantdn, 25 Mar 1978); 392 
(2900m, Gatero, 30 Apr 1978); 541 (2850m, Gigantdn, 13 Jan 1980); 638 (2888m, Carnacha, 
4 Mar 1 980); 891 (2870m, Carnacha, 4 Mar 1 978); 1 1 09 (2860 m, Mayhuayqui, 1 May 1 981 ). 
Cheilanthes myriophylla Desv. 

This species is distributed from Mexico and Central America to Chile, Brazil, and Argentina. 
In Peru it occurs on the western side of the Andes, and in inter-andean valleys, between 1 500m 



220 FERN GAZETTE: VOLUME 1 3 PART 4 (1 988) 



and 3500m elevation. In the study area it has been found in rock crevices in open areas. 
N. V. & I. F. 361 (2900m, Gigantdn, 26 Mar 1 978); 999 (2250m, Tarita, 21 Jan 1 981 ). 
Cheilanthes orbignyana Kuhn 

This species has been found in Peru and Bolivia. In the study area it grows in rock crevices 
and protected sites, together with Adiantum digitatum. 

N. V. & I. F. 534 (2860m, Gatero, 13 Jan 1980); 666 (2860m, Gatero, 4 Mar 1980); 1274 
(2860m, Gatero, 1 May 1981). 
Cheilanthes peruviana (Desv.) Moore 

This species occurs only in Peru, between 400m and 3200m elevation. It grows in rock 
crevices of open sites, together with Notholaena nivea and Pellaea ovata. 

N. V. & I. F. 352 (2860m, Gatero, 14 Jan 1980); 506 (2850m, Gigantdn, 13 Jan 1980); 642 
(2880m, Carnacha, 4 Mar 1980); 1276 (2870m, Gatero, 2 May 1981). 
Cheilanthes pilosa Goldm. 

This species occurs from Peru to Argentina. In Peru it occurs on the western side of the 
Andes and in inter-andean valleys, between 2800m and 4000m elevation. In the study area 
it grows in rock crevices and open sites. 
N. V. & I. F. 1 1 1 1 (2870m, Mayhuayqui, 3 May 1 981 ). 
Cheilanthes pruinata Kaulf. 

This species is distributed from Peru to Argentina. In Peru it occurs on the western side 
of the Andes and in inter-andean valleys, between 1500m and 4200m elevation. In the study 
area it has been found in rock crevices. 

N. V. & I. F. 345 (3000m, Carnacha, 25 Mar 1978); 533 (2860m, Gatero, 13 Jan 1980). 
Cystopteris fragilis(L.) Bernh. 

This is a cosmopolitan species. In Peru it has been found between 2400m and 4400m elevation. 
In the study area it grows in humid, shaded sites in the forest. 

N. V. & I. F. 277 (2850m, Gigantdn, 25 Mar 1978); 607 (2880m, Carnacha, 4 Mar 1980); 
724 (3300m, Ventanilla, 28 Jul 1 980). 
Elaphoglossum minutum (Pohl ex Fee) Moore 

This species grows in the Andes and southern Brazil. In the study area it has been found 
in rock crevices, protected by vegetation. 
B. L et al. 907 (2860m, Gigantdn, 8 Apr 1 986). 
Equisetum bogotense H.B.K. 

This species occurs from central America and the Greater Antilles to Chile and Argentina. 
In Peru it has been found between 500m and 4000m elevation. In the study area it grows 
commonly in humid places along running water. 

N. V. & I. F. 1 74 (2850m, Gigantdn, 28 Aug 1 977); 614 (2880m, Carnacha, 4 Mar 1 980). 
Notholaena nivea (Poiret) Desv. var. tenera (Hooker) Grisebach 

This species is distributed from Peru to Argentina. In Peru it occurs between 1800m and 
4000m elevation. In the study area it has been found in crevices in open areas together with 
Cheilanthes peruviana and Pellaea ovata. 
B. L. et al. 900 (2000m, Chunaca, 6 Apr 1 986). 
Pellaea ovata (Desv.) Weath. 

This species is distributed from Mexico to Argentina. In Peru it occurs in inter-andean valleys. 
It has been found in rock crevices of the lower part of the study area mixed with Cheilanthes 
peruviana and Notholaena nivea. 
B. L. etal. 901 (2000m, Chunaca, 6 Apr 1986). 
Pellaea ternifolia (Cav.) Link 

This species is distributed from southeastern U.S.A. to Argentina and the Hawaiian Islands. 
In Peru it occurs between 1600m and 4100m elevation. In the study area it grows in rock 
crevices in open sites. 

N. V. & I. F. 531 (2860m, Gatero, 1 3 Jan 1 980); 1110 (2880m, Mayhuayqui, 2 May 1 981 ). 
Polypodium pycnocarpum C. Chr. 

This species has been found in Peru and Bolivia. In Peru it occurs from La Libertad and 
Ancash to Puno and Cuzco, in the seasonal coastal vegetation ("lomas") and inter-andean valleys, 
between 350m and 4080m elevation. In the study area it grows as an epiphyte or in rock 
crevices in vegetation-protected sites. 

N. V. & I. F. 275 (2850m, Gigantdn, 25 Mar 1978); 328 (3000m, Gigantdn, 26 Mar 1978); 
526 (3000m, Pampa Zarate, 1 3 Jan 1 980). 
Selaginella novae-hollandiae (Sw.) Spring 

This species is distributed from Nicaragua to Bolivia. In Peru, it occurs between 350m and 
3200m elevation. In the study area, it grows in humid, protected sites. 



PTERIDOPHYTES OF ZARATE 221 



B. L. et al. 902 (2500m, Pascana, 6 Apr 1 986). 
Thelypteris glandulosolanosa (C. Chr.) R. Tryon 

This species is distributed in Ecuador, Peru and Bolivia. In Peru it has been found in Junin 
and Lima. In the study area, it grows in exposed sites in the forest. 

N. V. & I. F. 179 (2880m, Gigantdn, 30 Aug 1977); 515 (2850m, Gigantdn, Gigantdn, 13 Jan 
1 980); 670 (2825m, Gigantdn, 4 Mar 1 980). 
Thelypteris rufa (Poir.) A. R. Smith 

This species occurs from Ecuador to Peru and Bolivia. In Peru it has been found in the 
departments of Cajamarca, Lima, and Cuzco. In the study area it grows in humid sites protected 
by the vegetation. 

N. V. & I. F. 890 (3300m, Ventanilla, 28 Jul 1 980). 
Trachypteris induta (Maxon) R. Tryon & A. Tryon 

This species is endemic to Peru. It has been found in the departments of Amazonas, Cajamarca, 
La Libertad, and Lima between 750m and 2900m elevation. In the study area, which is close 
to the type locality, it grows in open sites in crevices. 

N. V. & I. F. 559 (2330m, Molle, 4 Mar 1980); 971 (2100m, Mital, 18 Jan 1981). 
Woodsia montevidensis (Spreng.) Hieron. 

This species is widely distributed in South America, from Colombia to Argentina and Uruguay. 
In Peru it occurs in inter-andean valleys and on both sides of the Andes above 2500m elevation. 
In the study area it grows in rock crevices partially protected by the vegetation. 
N. V. & I. F. 277 (2850m, Gigantdn, 25 Mar 1978); 344 (3000m, Carnacha, 25 Mar 1978); 
568 (2830m, Gigantdn, 4 Mar 1980); 638 (2880m, Carnacha, 4 Mar 1980); 1112 (2870m, 
Mayhuayqui, 1 May 1 981 ); 1 1 40 (2500m, Pascana, 1 May 1 981 ). 

DISTRIBUTION IN THE STUDY AREA 
In general, the number of pteridophyte species increases with elevation in the study 
area, although the species composition in any site varies with local topography and 
microhabitats. At lower elevations (1 500-2350m) only five species are present, while 
20 species occur in the forest (3000m). This difference is probably due to increased 
humidity with altitude, and the increased number of microhabitats. The forest only 
covers 1 5% (1 .5km 2 ) of the study area, but contains 87% of the species of pteridophytes. 

HABITATS 
The habitats where the pteridophytes occur are referred to using the classification 
of vegetation previously given by Valencia & Franke (1 980). The study area represents 
a good example of the variation in topography, altitude, and environments of the 
western Andes of central Peru. 

Cactus Zone (1 500-1 950m). This xerophytic vegetation is characterized by columnar 
cacti (Neoraimondia rosaiflora (Werd. & Backbg.) Backbg., Haageocereus acranthus 
(Vpl.) Backbg.), and some seasonal forbs (Jungia spp., Nicotiana spp., Lycopersicon 
peruvianum (L.) Mill.) or cultivated fields of Opuntia ficus-indica Mill. Pteridophytes 
do not occur here. 

Carica and Jatropha Zone (c. 1 950-2350m). This is an association of scattered shrubs 
of Carica candicansA. Gray and Jatropha macrantha Muell. Arg., and a larger number 
of seasonal forbs (Browallia americana L., Jungia spp., Lycopersicon peruvianum (L) 
Mill., Nicotiana spp., Piqueria peruviana (J.F. Gmel.) Robinson). In this zone we found 
Adiantum subvolubile, Cheilanthes myriophylla, C. peruviana, Notholaena nivea, and 
Pellaea ovata. 

Croton Zone (c. 2350-2600m). This dry scrub vegetation is dominated by Croton 
ruizianus Muell. Arg. Randia boliviana Rusby and Cordia macrocephala (Desv.) H.B.K. 
are also important. In this zone we found the terrestrial pteridophytes Adiantum 
subvolubile, Cheilanthes myriophylla, C. peruviana, Selaginella novae-hollandiae, and 
Trachypteris induta. The only exception was Polypodium pyenocarpum, which grows 
on lichen/moss mats found on some of the boulders. 



222 FERN GAZETTE: VOLUME 1 3 PART 4 (1 988) 



Thorny Scrub Zone (c. 2600-2700m). This is a transition zone from scrub vegetation 
to the forest. The dominant taxa are Hesperomeles cuneata Lindl., Duranta 
pseudorepens Mold., and Barnadesia blakeana Ferreyra. The pteridophytes that occur 
here are Adiantum digitatum, A. subvolubile, Cheilanthes myriophylla, C. peruviana, 
Polypodium pycnocarpum, Selaginella novae-hollandiae, and Woodsia montevidensis. 

Oligotherme forest (c. 2700-3200m). This is floristically the richest zone in the area, 
consisting of three dominant tree species, Oreopanax oroyanus Harms., Myrcianthes 
quinqueloba McVaugh, and Prunus rigida Koehne, along with scattered evergreen 
shrubs and several seasonal herbs. Here we found Adiantum digitatum, A. poiretii, 
A. subvolubile, Asplenium fragile, A. aethiopicum, A. sessilifolium, Cheilanthes 
myriophylla, C orbignyana, C. peruviana, C. pilosa, C. pruinata, Cystopteris fragilis, 
Elaphoglossum minutum, Equisetum bogotense, Pellaea ternifolia, Polypodium 
pycnocarpum, Selaginella novae-hollandiae, Thelypteris glandulosolanosa, T. rufa, and 
Woodsia montevidensis. 

MICPOHABITATS 

Four main microhabitats could be distinguished in the study area: crevices in open 
areas, sites protected by the vegetation, lichen/moss mats, and humid sites. These 
microhabitats are dispersed mosaic fashion, in the habitats previously described. The 
Carica and Jatropha zone had only two microhabitats, the Croton and Thorny Scrub 
zones had three, and the Oligotherme zone had all four microhabitats. 

The most important pteridophyte microhabitat was found in areas protected by 
the vegetation. We observed that the leaves and branches of shrubs and trees offer 
shade and protection at their base, keeping soil and air humidity high, and maintaining 
moderate temperatures. The pteridophytes which occurred here were Adiantum 
digitatum, A. poiretii, A. subvolubile, Asplenium fragile, Asplenium sessilifolium, 
Cheilanthes orbignyana, Cystopteris fragilis, and Elaphoglossum minutum. In the lower 
and drier zones of the study area, this microhabitat also permits pteridophytes, such 
as Selaginella novae-hollandiae, to grow together with drought-evading forbs and 
graminoids. 

Crevices in open areas also represent an important microhabitat; they occur in 
rocky places or cliffs. This microhabitat probably provides soil and nutrients, and a 
partial protection from insolation. This type of microhabitat is especially frequent in 
the lower parts of the study area. Most of the cheilanthoid species, which were 
described by Mickel (1986) as generally occurring in dry, rocky areas, were well 
represented here: Cheilanthes myriophylla, C. peruviana, C. pilosa, C pruinata, 
Notholaena nivea, Pellaea ovata, P. ternifolia, and Trachypteris induta. In addition, 
Asplenium aethiopium and Woodsia montevidensis also ocurred here. 

The lichen/moss mat microsite was found on tree branches or boulders in the 
forest and nearby zones. The only pteridophyte in this microhabitat was Polypodium 
pycnocarpum, many individuals of which grew here as epiphytes. 

The fourth microhabitat consisted of humid sites. These mainly occur in very humid 
rocky places and beside running water, in the upper part of the forest. Equisetum 
bogotense, Thelypteris glandulosolanosa, and T. rufa were found here surrounded 
by forbs. 

BIOGEOGRAPHICAL ELEMENTS 

We distinguish four groups of species in the study area, on the basis of types of 

distribution ranges: tropical-subtropical, tropical American, Andean, and cosmopolitan. 

The tropical-subtropical group includes those species occurring in the tropics and 

subtropics of America, and elsewhere. This group contains 12 species: Adiantum 



PTERIDOPHYTES OF ZARATE 223 



poiretii, Asplenium aethiopicum, A. fragile, A. sessilifolium, Cheilanthes myriophylla, 
C. pilosa, C. pruinata, Equisetum bogotense, Notholaena nivea, Pellaea ovata, P. 
ternifolia, and Woodsia montevidensis. The tropical American group includes those 
species restricted to the tropics of America. It contains only one species: Selaginella 
novae-hollandiae. The Andean group is restricted to species that occur along the 
Andes and contains nine species, including two endemic to Peru: Adiantum digitatum, 
A. subvolubile, Cheilanthes orbignyana, C. peruviana, Elaphoglossum minutum, 
Polypodium pycnocarpum, Thelypteris glandulosolanosa, T. rufa, and Trachypteris 
induta. The cosmopolitan group contains only Cystopteris fragilis. 

The forest of the study area contained all the biogeographical groups, in addition 
to the largest number of species and of microhabitats. By including it and the 
surrounding area within a nature reserve, a valuable portion of the pteridophyte species 
of the western central Andes would be preserved, thus leaving open the possibility 
in the future of achieving a better understanding of the history and development 
of the flora of the western Andes. 

ACKNOWLEDGEMENTS 
We are very grateful to K. Young, who discussed and corrected the manuscript. Also, 
we thank R. G. Stolze for reading the manuscript, and M. Lane for facilities during 
preparation of this article. Special thanks go to our friends I. Franke and A. Cano 
for help in the field and comments. We are grateful to the kind people of the town 
of San Bartolome for their assistance. The field work was supported by the Frankfurt 
Zoological Society (project 905/81 ) and the Consejo Nacional de Ciencia y Tecnologfa 
(CONCYTEC)of Peru. 

REFERENCES 
CHAPMAN, F.M. 1926. The distribution of Bird-life in Ecuador. A contribution to a study of 

the origin of Andean bird-life. Bull. Amer. Mus. Nat. Hist. 55: 1-784. 
FRANKE, I. & VALENCIA, N. 1984. Zarate: Una unidad de Conservation. Report from the Museo 

de Historia Natural "Javier Prado". Lima. 
KOEPCKE, H. 1961. Synekblogische Studien an der Westseite der peruanischen Anden. Bonn 

Zool. Beitr. 29: 1 -320. 
KOEPCKE, H. W. & KOEPCKE, M. 1958. Los restos de bosques en las Vertientes Occidentales 

de los Andes Peruanos. Bol. Comite Nac. Protection de la Naturaleza, Lima 16: 22-30. 
KOEPCKE, M. 1954. Corte ecoldgico transversal en los Andes del Peru Central, con especial 

consideracibn de las Aves. I. Costa, Vertientes Occidentales y Region Altoandina. Mem. 

Mus. Hist. Nat. "Javier Prado" 3: 1 -1 1 9. 
KOEPCKE, M. 1958. Die Vogel des Waldes von Zarate (Westhang der Anden in Mittel Peru). 

Bonn Zool. Beitr. 9: 130-193. 
LEON, B. 1983. Los Pteridofitos de la ciudad de Lima y alrededores. Unpublished Lie. thesis, 

Universidad Nacional Mayor de San Marcos. Lima, Peru. 
MICKEL, J. T. 1986. Ecology and geography of the cheilanthoid-gymnogrammoid ferns. Amer. 

J. Bot. 73:731. 
SANCHEZ, I. 1965. Las Polypodiaceas de la Provincia de Trujillo. Unpublished Br. thesis, 

Universidad Nacional de Trujillo. Trujillo, Peru. 
TRYON, R. 1 960. The ecology of Peruvian ferns. Amer. Fern J. 50: 46-55. 
TRYON, R. 1964. The ferns of Peru. Polypodiaceae (Dennstaedtiaeae to Oleandreae). Contr. Gray 

Herb. 97:1-253. 
TRYON, R. 1 971 . Ferns of the Andes and Amazon. Morris Arbor. Bull. 22: 7-13. 
VALENCIA, N. & FRANKE, I. 1980. El bosque de Zarate y su conservacidn. Bol. de Lima 7: 

76-86; 8: 26-35. 
VASQUEZ, L. P. 1967. Las Polypodiaceas de la Provincia de Contumaza. Unpublished Br. thesis, 

Universidad Nacional de Trujillo. Trujillo, Peru. 
WEBERBAUER, A. 1936. Phytogeography of the Peruvian Andes. In Macbride, J.F. (Ed.) "Flora 

of Peru", Field Mus. Nat. Hist. Bot. ser. 13: 13-80. 
WEBERBAUER, A. 1945. El Mundo Vegetal de los Andes Peruanos. Ministerio de Argricultura, 

Lima. 



224 FERN GAZETTE: VOLUME 1 3 PART 4 (1 988) 

APPENDIX 1 

KEYTOTHETAXA 
A Axes or stems without conspicuous leaves, these reduced to scales in a whorled 

arrangement Equisetum bogotense 

A' Axes or stems with conspicuous leaves B 

B Axes prostrate; leaves in four longitudinal rows Selaginella novae-hollandiae 

B' Axes erect, plants not as above C 

C Lamina entire or pinnatisect D 

C Lamina pinnate or pinnate-pinnatifid F 

D Lamina entire, lanceolate Elaphoglossum minutum 

D' Lamina pinnatisect E 

E Petiole articulate; lamina lanceolate; lamina scales brown or gold-brown, somewhat dense 

on the abaxial surface Polypodium pycnocarpum 

E' Petiole not articulate; lamina pedate or 3-lobed; lamina scales pinkish brown, sparse on 

the abaxial surface Trachypteris induta 

F Lamina with last segments of the central pinna less than 5mm long; last segments elliptic, 

orbicular or deltoid G 

F' Lamina with last pinnae or segments of the cental pinnae more than 5mm; last segments 

cuneate, flabellate or elliptic L 

G Lamina with scales of trichomes; when glabrous then herbaceous (Cheilanthes 

orbignyana) H 

G' Lamina glabrous, chartaceous Notholaena n/Vea var. tenera 

H Leaves scaly I 

H' Leaves glabrescent or pubescent J 

I Lamina usually 3-4 pinnate; segments suborbicular, less than 1 mm long 

Cheilanthes myriophylla 

V Lamina 1 -2 pinnate-pinnatifid; segments triangular, more than 1 mm long 

Cheilanthes peruviana 

J Lamina herbaceous, glabrescent Cheilanthes orbignyana 

J' Lamina chartaceous or coriaceous, pubescent K 

K Axes with non-glandular trichomes Cheilanthes pilosa 

K' Axes with glandular trichomes Cheilanthes pruinata 

L Lamina with 3-4 pairs of gradually reduced basal pinnae, lamina more than 5cm 

wide , M 

L' Lamina generally without reduced basal pinnae, lamina less than 5cm wide N 

M Lamina herbaceous-chartaceous; central pinnae less than 10cm long; leaves 60cm long; 

indusia inconspicuous Thelypteris rufa 

M' Lamina coriaceous; centrai pinnae 10-1 5cm long; leaves 100cm long; indusia conspicuous, 

villous Thelypteris glandulosolanosa 

N Lamina pinnate-pinnatifid, if bipinnate then chartaceous 

N' Lamina 2-4 pinnate, herbaceous T 

O Segment margins entire P 

0' Segment margins crenate or partially dentate Q 

P Axes stramineous; pinnae petiolulate Pellaea ovata 

P' Axes dark castaneous or atropurpureous; pinnae not petiolulate Pellaea ternifolia 

Q Indusium attached by one side to the segment; pinnae bases cuneate R 

Q' Indusium cup-like; pinnae base not as above Woodsia montevidensis 

R Lamina chartaceous, scaly; lamina scales subulate and conspicuous 

Asplenium aethiopicum 

R' Lamina herbaceous, glabrescent S 

S Pinnae less than 1cm long, pinnae entire or with an acroscopic lobe 

Asplenium fragile 

Pinnae more than 1cm long, generally lobed in the acroscopic and basiscopic 

side Asplenium sessilifolium 

T All ultimate segments free, cuneate or flabellate; marginal false indusium U 

T' Only some ultimate segments free, cuneate; indusium scale-like Cystopteris fragilis 

U Segments articulate; leaves sub-scandent Adiantum digitatum 

U' Segments not articulate; leaves erect V 

V Petiolule of the first acroscopic pinnule of the basal pinna less than 2mm long; distance 
between the first pinnule and the rachis less than 3mm Adiantum subvolubile 

V Petiolule of the first acroscopic pinnule of the basal pinna more than 2mm long; distance 
between the first pinnule and the rachis more than 3mm W 

W Ceraceous yellow glands absent or restricted among the sporangia 

Adiantum poiretii var. poiretii 

W" Ceraceous yellow glands on the abaxial segment surface 

Adiantum poiretii var. sulphureum 



FERN GAZ. 13(4)1988 225 



A FIELD SURVEY OF PTERIDIUM AQUILINUM 
(DENNSTAEDTIACEAE: PTERIDOPHYTA) MYCORRHIZAS 



H.M. JONES 1 and E. SHEFFIELD 2 

Department of Cell & Structural Biology, University of Manchester 

Williamson Building, Manchester M13 9PL 

^Present address: Department of Botany, University of Sheffield, Western Bank, 

Sheffield S10 2TN. 
2 To whom correspondence should be addressed. 

ABSTRACT 
Roots of Pteridium aquilinum were sampled from natural populations in the Northern 
Hemisphere and routinely found to contain mycorrhizal fungi. Amount of infection 
varied greatly between sites, but was not correlated with soil moisture, pH, nitrogen 
or phosphorus content. The significance of these findings in relation to Pteridium 
grown in controlled conditions is discussed. 

INTRODUCTION 
It is clear that very few plants growing in natural plant communities are free from 
mycorrhizal infection, and studies which have included pteridophytes indicate that 
they are no exception (e.g. Lohman 1927; Bouillard 1957). Fungal enhancement of 
phosphorus (P) uptake has been found to be the main reason for improved growth 
in infected plants and the uptake of other elements and water may also be important 
(see Read 1984, for review). Experiments with Southern Hemisphere Pteridium 
aquilinum (L.) Kuhn (subsp. caudatum (L.) Bonaparte var. esculentum (Forster) Kuhn) 
have shown that mycotrophic plants grow significantly better than uninfected controls 
in soils with low P levels (Cooper 1975) and that roots of Pteridium growing in New 
Zealand are consistently mycorrhizal (Cooper 1976). Cooper concluded that the 
available P in the soil influenced the extent to which roots became mycorrhizal (Cooper 
1976). There are reports of Pteridium aquilinum (subsp. aquilinum var. aquilinum) 
mycorrhizas in Britain (e.g. Conway & Arbuthnot 1949; Hepden 1960) but these refer 
to very few plants, sites and soil types. Pteridium frequently inhabits nutrient-poor 
soils in Britain but the role played by mycorrhizal infection in the widespread success 
of this species has never been assessed. The aim of the present study was to investigate 
the incidence of mycorrhizas in well-established Northern hemisphere Pteridium 
populations and to relate it to soil parameters. 

MATERIALS AND METHODS 
Twelve well-established Pteridium sites were sampled during the summer months: 
Stiperstones, Shropshire (SO 372976); White Nancy, Bollington, Cheshire (SJ 939772); 
Winterside Farm, Bollington, Cheshire (SJ 955780); Bakestonedale Moor, Pott Shrigley, 
Cheshire (SJ 955798); Birch Knoll, Langley, Cheshire (SJ 932722); Ladybower, 
Bamford, Derbyshire (SK 202855, 21 7876); Tegsnose Country Park, Langley, Cheshire 
(SJ 948723); Erwood Hall, Derbyshire (SK 004754); Buddock, Cornwall (SW 786324); 
Nr. Owslebury, Hampshire (SU 473234); Richmond Park, Surrey (TQ 207742); Nr. 
Harleston, Norfolk (TM 250835). 

Rhizomes and roots were excavated and treated as follows. The dry weight of 
rhizomes and roots in 1m x 1m x 15cm deep areas were determined and infection 
levels of sub-samples of roots were estimated. Roots were also excavated from leading 
edges and central portions of well-established stands. Infection levels were 



226 FERN GAZETTE: VOLUME 13 PART 4 (1988) 



microscopically assessed in 1cm segments of roots which had been washed, cleared 
in 10% KOH, acidified and stained in lactophenol blue according to the method of 
Phillips & Hayman (1970). Roots which could not be processed immediately were 
fixed in FAA, those which did not clear after several hours at 90°C in KOH were 
cleared with hydrogen peroxide. Treatment with alkaline hydrogen peroxide (after 
Philips & Hayman) failed to clear Pteridium roots, but freshly made (acidic) 10 vols 
solutions of hydrogen peroxide rapidly cleared them. Pilot experiments in which 
infection was recorded as absence or presence of infection per root segment indicated 
that this method gave less accurate results than percentage root length infected (after 
Biermann & Linderman 1981). The latter method was therefore adopted throughout 
the study: between 40 and 150 segments of root per sample were examined, the 
percentage length infected estimated, a mean value calculated for each sample and 
the 95% confidence limits and the standard errors calculated. 

Fresh roots from the Cornish sample were also attached to a scanning electron 
microscope (SEM) slug and plunged into slushed liquid nitrogen inside a Hexland 
slushing chamber. The chamber was evacuated and the slug transferred to a Cambridge 
S150 SEM fitted with a (Hexland) stage cooled with nitrogen at -190°C. The roots 
were sliced open with a cooled blade and transferred into the SEM. The specimens 
were briefly warmed to -80°C to remove.ice, withdrawn into a pre-chamber at -180°C 
and sputter-coated with gold. The coated roots were then returned to the SEM at 
-180°C and photographed. 

Additional Pteridium sites in Switzerland, Spain, Hungary, NE and NW USA were 
also sampled and analysed for presence and absence of mycorrhizal infection. 

Soil samples were taken from around the rhizomes at each site and analysed. 
Soil pH was measured using the method of Peech (1965). Total nitrogen and 
phosphorus, and available phosphorus content were measured using the methods 
of Black et at. (1 965). Soil moisture content was assessed by oven drying the samples 
for 18 hours at 105°C and subtracting the dry from the wet weight. Results were 
adjusted to reflect amounts in grams of dry soil. 

RESULTS 

Microscopic analysis 

The roots of all the specimens examined showed evidence of mycorrhizal infection. 
The majority of roots contained fungal elements which stained blue with lactophenol 
blue within the tissues, a small number contained, in addition, limited areas occupied 
by yellow/brown coloured fungal structures. As the blue-staining mycorrhizal infection 
was by far the most common and widespread only this will be described in detail. 

In heavily infected specimens, a continuous layer occupied by fungi occurred 
throughout many roots. This layer consisted of a fungal zone, one or two cells deep, 
immediately outside the stele, e.g. Fig. 1a and b. Infection was never observed within 
the stele or meristematic tissues, and was never associated with signs of tissue damage 
or necrosis. The mycorrhizal layer was interrupted at sites of lateral root emergence. 
Infected cells contained either finely divided arbuscules and hyphae (e.g. Fig. 1c) or 
coils of coarser hyphae (e.g. Fig. 1 d). Some specimens contained thick-walled spherical 
vesicles throughout the cortex {e.g. Fig. 1e). In roots with low levels of infection, 
fungal structures were often limited to discrete patches of inner cortex and such 
limited areas were often connected to a single external hypha. External hyphae were 
never found to have formed ectomycorrhizal sheaths but penetrated roots directly 
or, where root hairs were present, by entering the distal part of the hair. Once in 
the outer cell layers the hyphae branched, giving rise to hyphae which penetrated 



PTERIDIUM AQUILINUM MYCORRHIZAS 



227 




Figure 1. All photo-micrographs show Pteridium aquilinum mycorrhizal infection, a) Cryo SEM 
of fractured root to show cells outside the stele occupied by fungal structures x 100; one cell 
enlarged in b), x 400. c) - e) Light micrographs of cleared, stained roots, c) Finely divided arbuscule, 
x 300. d) Hyphal coils, x 300. e) Spherical vesicles and hyphae, x 360. 



228 



FERN GAZETTE: VOLUME 13 PART 4 (1988) 



cortical cells next to the stele. Hyphae also ran through the cortex, parallel to the 
long axis of the root, and penetrated inner cortical cells to either side of the original 
point of entry into the root. 

Site analysis 

Soil moisture ranged from 0.22-3.72g H 2 (g dry soil -1 ), total nitrogen from 2.09- 
12.48umol, total phosphorus from 1 09-5.79>umol, available phosphorus from 0.02- 
1.81, and soil pH from 2.4-5.3. The infection levels of roots sampled from leading 
edges of Pteridium and those from the centre of well established stands did not differ 
significantly. Although infection levels differed significantly between sites, the 
differences were not correlated with differences in soil moisture, total nitrogen, total 
or available phosphorus content, soil pH, or dry weight of rhizomes and roots 
nrr . As phosphorus was thought to be the most important site variable in the present 
context Fig. 2 is included to illustrate the data relating to available phosphorus and 
amount of infection. 



"O 60 
CD 

O 
CD 

S 50 



c 



d^ 

C 
CD 
CD 



40 



k'l. 



5 1.0 1.5 2.0 

Available Phosphorus (/am g dry soil 



Figure 2. Mean percentage length of mycorrhizal infection in Pteridium aquilinum root segments 
plotted against available phosphorus in soil from nineteen different sites, mean percentage length 
infected was calculated according to the method of Biermann & Linderman (1981) and bars 
show the standard error about each mean. Available phosphorus was calculated according to 
the method of Black et al. (1 965). 



PTERIDIUM AQUILINUM MYCORRHIZAS 229 



DISCUSSION 

The results of the present study do not confirm the contention of Cooper (1976) that 
the available P in the soil influences the extent to which Pteridium roots become 
mycorrhizal. This lack of agreement is apparent rather than real, however, as Cooper 
based her statement on the results of uninfected versus infected plants grown in 
pots of soil with various P levels (Cooper 1975). In these experiments there was 
a clear influence of P level on percentage infection. In the field Cooper herself reported 
that root samples from soils relatively high in available P seemed to be no different 
from the rest (Cooper 1 976, see also Hepden 1 960). In this respect, therefore, Pteridium 
in the Northern Hemisphere resembles Pteridium in the Southern Hemisphere, but 
the reasons for the difference between experimental and field data are not immediately 
apparent. From Cooper's (1975) results a lower level of infection would be expected 
in plants growing in soils of high P. If plants routinely become infected, perhaps 
even at the prothallial stage, it may be that those growing in poor soils derive more 
benefit from the association than those in rich soils. Since soil pH, nitrogen, phosphorus 
and moisture levels are not correlated with infection, the present study provides no 
indication of factors which determine levels of infection. Higher or lower levels of 
infection must reflect site parameters untested in this investigation, or individual 
differences between plants. Pteridium in Britain is well known to be highly polymorphic 
(e.g. for cyanogenesis, Hadfield & Dyer 1986; in DNA composition, Jubrael, Sheffield 
& Moore 1986; and in morphology, Page 1986) so levels of infection may reflect 
another inherent polymorphism, perhaps linked with factors which account for 
variations observed in susceptibility to pathogenic fungal infection (Burge pers. comm). 
Although this study cannot account for the differences found in infection levels 
it is clear that Northern Hemisphere Pteridium is routinely mycorrhizal. This has 
important implications for studies of mineral nutrition of the fern, and casts a new 
light on those which have, in the past, been conducted with plants raised from spores 
in axenic conditions. Where such plants have been cultivated in sterile media (e.g. 
Schwabe 1 951 , 1 953; Conway & Stephens 1 956) they cannot have been mycorrhizal, 
so the results of these studies, upon which much of the recent work on nutrient 
cycling has been based, are questionable. Definitive studies of Pteridium nutrient 
cycling clearly require natural, not sterile, conditions, and it seems likely that this 
applies to most ferns, in view of the large body of data on fern mycorrhizas (e.g. 
Lohman 1927; Hepden 1960; Bouillard 1957). Fungal symbionts do not seem to have 
figured in plans to combat the "bracken problem" and it is feasible that Pteridium 
on poor soils would be severely disadvantaged by fungicides. Since experimental plants 
are known to grow very poorly in the absence of mycorrhizal fungi in soils of low 
P (Cooper 1975), elimination of the fungal component might provide a new control 
strategy. Fungicides could be combined with herbicides to provide control in high 
P soils. 



ACKNOWLEDGEMENTS 

We are grateful to the Trustees of the Nuffield Foundation for financial support, to 
Ms S.M. Crowe for excellent technical assistance, and to Mr J.H. Bastin, Mr B. Burrows, 
Mr N. and J. Barber, Dr J.J. Schneller, Prof. PR. Bell, Dr B.J. Way, Dr R.J., Mr W. 
and Mrs J.A. Drakeley, Dr M.G. and Mrs V. Gregg, Prof. G. Vida, Mr P.G. Wolf and 
Dr D.S. Conant for help with collection of material. 



230 FERN GAZETTE: VOLUME 1 3 PART 4 (1 988) 



REFERENCES 

BIERMANN, B & LINDERMAN, R.G. 1981. Quantifying vesicular-arbuscular mycorrhizae: a 

proposed method towards standardization. New Phytol. 87: 63-67. 
BLACK, C.A., EVANS, D.D., WHITE, J.L, ENSMINGER, LE. and CLARK, F.E. 1965. Methods 

of soil analysis. Am. Soc. Agr. Wisconsin, USA. 1572 pp. 
BOUILLARD, B. 1957. La mycotrophie chez les pteridophytes. Sa frequence, ses caracteres, 

signification. Le Botaniste 41 : 5-185. 
CONWAY, E. & ARBUTHNOT, M. 1949. Occurrence of endotrophic mycorrhiza in the roots of 

Pteridium aquilinum Kuhn. Nature 163: 609-610. 
CONWAY, E. & STEPHENS, R. 1956. Sporeling establishment in Pteridium aquilinum: Effects 

of mineral nutrients. J. Ecol. 45:389-399. 
COOPER, K.M. 1975. Growth responses in Solanum, Leptospermum and New Zealand ferns. 

In F.E. Sanders, B. Mosse and P.B. Tinker (eds) Endomycorrhizas. Academic Press, London: 

391-407. 
COOPER, K.M. 1976. A field survey of mycorrhizas in New Zealand ferns. N.Z. J. Bot. 14: 169- 

181. 
HADFIELD, P.R. & DYER, A.F. 1986. Polymorphism of cyanogenesis in British populations of 

bracken (Pteridium aquilinum (L.) Kuhn). In R.T. Smith and J.A. Taylor (eds) Bracken; 

ecology, land use and control technology: 293-301 . 
HEPDEN, P.M. 1960. Studies in vesicular-arbuscular endophytes. II. Endophytes in the 

Pteridophyta, with special reference to leptosporangiate ferns. Trans. Brit. Myc. Soc. 43: 

559-570. 
JUBRAEL, J. M.S., SHEFFIELD, E. & MOORE, D. 1986. Polymorphisms in DNA of Pteridium 

aquilinum. In R.T. Smith and J.A. Taylor (eds) Bracken; ecology, land use and control 

technology. 309-315. 
LOHMAN, M.L 1927. Occurrence of mycorrhiza in Iowa forest plants. Stud. Nat. Hist, la Univ. 

11:5-30. 
PAGE, C.N. 1986. The strategies of bracken as a permanent ecological opportunist. In R.T. Smith 

and J.A. Taylor (eds) Bracken; ecology, land use and control technology; 1 73-1 83. 
PEECH, M. 1965. Hydrogen Ion Activity. In: Methods of soil analysis. C.A. Black (Chief Ed). 

Am. Soc. Agr. Wisconsin, U.S.A. 
PHILLIPS, J.M. and HAYMAN, D.S. 1970. Improved procedures for clearing roots and staining 

parasitic and vesicular-arbuscular mycorrhizal fungi for rapid assessment of infection. 

Trans. Br. Myc. Soc. 55: 1 58-1 61 . 
READ, D.J. 1984. The structure and function of the vegetative mycelium of mycorrhzal roots. 

In The ecology and physiology of the fungal mycelium. Brit. Myc. Soc. Symp. 8, CUP: 

215-240. 
SCHWABE, W.W. 1951. Physiological studies in plant nutrition XVI. The mineral nutrition of 

bracken. Part I. Prothallial culture and the effects of phosphorus and potassium supply 

on leaf production in the sporophyte. Ann. Bot. (NS) 15: 41 7-446. 
SCHWABE, W.W. 1953. Physiological studies in plant nutrition XVI. The mineral nutrition of 

bracken. Part II. The effects of phosphorus and potassium supply on total dry weights, 

leaf areas, net assimilation rates, starch and water content etc. in the sporophyte. Ann. 

Bot. (NS) 1 7. 225-262. 






FERN GAZ. 13(4)1988 231 



ADAPTIVE STRATEGIES OF MARSILEA (MARSILEACEAE: 
PTERIDOPHYTA) IN THE LAKE CHAD BASIN OF N.E. NIGERIA 

JAN KORNAS 

Institute of Botany, Jagiellonian University, 

ul. Lubicz46, 31-512 Krakow, Poland 

ABSTRACT 

Five species of Marsilea: M. berhautii, M. distorta, M. minuta, M. nubica and M. 
subterranea occur together near Maiduguri in the Lake Chad Basin of N.E. Nigeria. 
They differ from one another in their habitat requirements (with regard to water 
and soil conditions), in their seasonal patterns of growth, reproduction and dormancy, 
as well as in the position of the sporocarps on the plant and the mode of their 
protection against adverse external influences. All these characters seem to be 
of a selective value and apparently originated through an adaptive radiation of 
sympatric taxa into various ecological niches. 

INTRODUCTION 
The genus Marsilea L. includes c. 50-70 species distributed in the tropical and warm 
temperate zones of all continents (Cook et al. 1974, Gupta 1962, Johnson 1986, 
Launert 1968, Sadebeck 1902, Tryon & Tryon 1982, Willis 1973). Distinct diversity 
centres are located in the semi-arid areas of Africa just on the borders of the deserts 
(Table 1): in the western and eastern parts of Sub-Saharan Africa (10 species and 
14 species, respectively), as well as in Southern Africa (16 species) (Launert 1968, 
1970, 1971, 1984, Schelpe & Anthony 1986). This unusual diversity apparently 
originated through an adaptive radiation of sympatric taxa into various ecological niches; 
an evolutionary process which most probably was stimulated by the extremes of the 
semi-arid climate, first of all the striking contrasts between life conditions in the 
rainy and the dry seasons (Kornas 1 985). 

The aim of the present paper is to illustrate this situation with data collected 
near Maiduguri in the Lake Chad Basin of N.E. Nigeria, where I had the opportunity 
to study the vegetation changes during two rainy seasons and the intervening dry 
season, in 1978 and 1979. The field work was partly sponsored by the University 
of Maiduguri, Nigeria. The final version of the paper was prepared under a grant 
of the Polish Academy of Sciences (project MR-ll.2.2.4). 

The specimens collected are deposited in the Herbarium Universitatis Jagiellonicae 
Cracoviensis (KRA), with duplicates in Kew (K) and the University of Maiduguri 
Herbarium. 

STUDY AREA 
The Lake Chad Basin of N.E. Nigeria is an extremely flat country with almost no 
local relief (Fig. 1). It occupies the former lake bottom of the Early Holocene, and 
it is limited in the south by the old shore line (Bama Ridge) at an altitude of c. 330m 
(Grove 1970). It is covered with a mosaic of recent deposits of the lake and its deltas, 
ranging from highly impermeable, black, cracking lagoonal clays to sands (Land Systems 
1970). The climate of the area is of an extremely hot semi-arid type (tropical climate 
with summer rains, type II of Walter 1971). The mean annual temperature exceeds 
26°C, the absolute maximum temperature reaches above 42°C, and the mean annual 
rainfall is about 600mm (Fig. 2). There is a short rainy season of c. 4 months (June- 
September) and a prolonged season of drought of c. 8 months (October-May). The 
plant cover consists mainly of thorn savannas of various types (Sudanian woodland 
and wooded grassland of White 1983), most of them being highly degraded by 
overgrazing and fuel wood collecting. 



232 



FERN GAZETTE: VOLUME 1 3 PART 4 (1 988) 



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gibba 

minuta var. minuta 

subterranea 

aegyptiaca 

coromandelina 

distorta 

nubica var. nubica 

nubica var. gymnocarpa 


aethiopica Launert 
botryocarpa F. Ballard 
fadeniana Launert 
megalomanica Launert 


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gibba A. Br. 

minuta L. var. minuta 

subterranea Lepr. ex A. Br 

aegyptiaca Willd. 

coromandelina Willd. 

distorta A. Br. 

nubica A. Br. var. nubica 

nubica A. Br. var. gymnocarpa 

(Leor. ex A. Br.) Launert 




^ S 


^^^^^2^2 




















MARSILEA IN NIGERIA 



233 




50 100 150 200 km 

1 1 I I I 



FIGURE 1 . Map of North-Eastern Nigeria. 

At the end of the rains vast areas of the savanna are covered with innumerable 
shallow pools, which shrink during the dry season and uncover their muddy bottoms. 
A rich ephemeral aquatic and amphibious vegetation appears in these pools. It is 
composed of a variety of short-lived species, either annuals or perennials, with their 
perennating organs buried in the mud. The angiosperms with deeply hidden corms, 
Aponogeton conjugatus Schum. & Thonn., Burnatia enneandra M. Micheli and 
Nymphaea micrantha Guill. & Perr., are good examples of this life-form (the 
nomenclature of the angiosperms follows Hutchinson & Dalziel 1 954-1 972). 

There is a considerable share of pteridophytes in the flora of seasonal pools in 
the Lake Chad Basin, including the quillwort Isoetes schweinfurthii A. Br. (with 
perennating corms) and five species of the water fern Marsilea (with two kinds of 
perennating organs: creeping rhizomes and hard, thick-walled sporocarps): M. berhautii 
(Fig. 3), M. distorta{F\g. 4), M. minuta(F\gs. 5, 6), M. nubica{F\g. 7) and M. subterranea 
(Fig. 8) (for a complete list of specimens collected see Kornas 1983). That so many 
Marsilea species are able to coexist in the study area is evidently because they occupy 
different ecological niches. 



ECOLOGICAL SPECIALIZATION IN MARSILEA 
The Marsilea species of the Lake Chad Basin differ remarkably from one another 
in their habitat requirements (with regard to the water and soil conditions), in their 
seasonal patterns of growth, reproduction and dormancy, as well as in the location 
of sporocarps on the plant and the mode of their protection against adverse external 
influences, especially desiccation. One species, M. berhautii, seems to be strictly 
aquatic, with the rhizomes rooting in the bottom of pools (up to 0.5m deep) and 
the stems and leaves floating on the water surface (Fig. 3). I did not see terrestrial 
forms of this species, neither did I find indications of such forms in the literature 



234 



FERN GAZETTE: VOLUME 1 3 PART 4 (1 988) 



MAIDUGURI 354m 26.5° 639 




M. nubica 
M. berhautii 

M. distorta 

M. minuta 

M. subterranea 



• • • • 



• 


• 

• •• 

• • • 


• • 
• • • 




• 
• 




• 
• 





VI 



XII II 



FIGURE 2 Climate diagram of Maiduguri drawn after Walter & Lieth 1 960 (above), and phenology 
of Marsilea species in the Lake Chad Basin near Maiduguri in 1978-1979 (below): each dot 
designates one herbarium collection. 



(see Johnson 1986: 13, Launert 1968: 279). On the contrary, M. distorta (Fig. 4) 
appears in pools usually only when they are already completely dry, and thus represents 
a truly terrestrial plant (see Launert 1968 285); its aquatic form with submerged 
rhizomes and floating leaves occurs only exceptionally (I collected it only once, in 
a pool 5 cm deep). The three remaining species, M. minuta, M. nubica and M. 
subterranea, are very typical amphiphytes: they start their development submerged 
in shallow water and finish it completely emerged on dry soil. The first, aquatic, 
phase of their growth is purely vegetative; the second, terrestrial, phase is reproductive, 
with a profuse formation of sporocarps (Figs. 5, 6. 7. 8). The transition from the 
aquatic phase to the terrestrial one is connected with far-reaching changes in the 
morphology and anatomy of the whole plant, including roots, rhizomes and fronds 
(as described for various Marsilea species e.g. by Allsopp 1 954 & 1 963. Gaudet 1 964, 
Gluck 1 922 & 1 936. Goebel 1 91 8, Gopal 1 968, Gupta 1 962. Sadebeck 1 902, Schmidt 
1978. White 1966 and others). The main differences between water forms and land 
forms of Marsilea species are indicated in Table 2 and illustrated in Figures 5-7. 



MARSILEA IN NIGERIA 



235 



TABLE 2 
Morphological differences between aquatic and terrestrial plants in amphiphytic Marsilea species 





Aquatic plants with floating leaves 


Terrestrial plants with aerial leaves 


Rhizome internodes 


elongated 


short 


Roots 


thick and relatively unbranched 


thin and repeatedly branched 


Petioles 


long and flaccid 


short and stiff 


Leaflets 


large 


small 


Terminal margins of leaflets 


entire 


crenulate or serrulate, or 

superficially to deeply lobed 


Reddish streaks on under surface 






of leaves 


often present 


absent 


Anatomical structure 


less highly differentiated 


more highly differentiated 


Hair-covering on vegetative parts 


few or no hairs 


copious hairs 


Aerenchymatic tissues in roots. 






rhizomes and petioles 


well developed 


much less developed 


Storoata 


only on the upper surface of leaves 


on both surfaces of leaves 




(epistomatic), not sunken 


(amphistomatic), sunken 


Leaf cuticle 


thin on the lower surface, thick 
on the upper surface 


thick on both surfaces 


Sporocarps 


usually absent 


usually present 



Edaphic requirements of Marsilea species in the Lake Chad Basin are clearly 
differentiated (similarly as e.g. in the Indian species - Gopal 1 969). M. distorta evidently 
requires light sandy soils. It seems that only on such soils this truly terrestrial plant 
is able to bury underground its growing sporocarps, even when the substratum is 
already dry. Three other species grow on heavy clay, either exclusively (M. subterranea) 
or nearly so (M. minuta, M. nubica). Such soils usually contain large amounts of 
organic matter. As demonstrated by Gopal (1 969) the organic matter in soil considerably 
promoted growth in the Indian species of Marsilea which he investigated. 

The phenological sequence of the five species occurring in the study area is quite 
distinct (Fig. 2). The most precocious among them is M. nubica, in which fully developed 
sporophytes with sporocarps could already be collected in September before the rains 
stopped. M. distorta was found from the very beginning of the dry season (in October), 
M. minuta a few weeks later (in November), while M. subterranea emerged most 
tardily (in January), when the rains were already over for a couple of months and 
even the most persistent pools were nearly dry. In places where two species of Marsilea 
occurred side by side, a concentric zonation was noticed, with the more precocious 
species on the periphery and the tardier one in the centre of the pool (Fig. 9). Under 
such circumstances the outer species may already appear in its land form and pass 
through its second, reproductive phase, with plenty of sporocarps, while the central 
one still remains in the first, vegetative phase and displays the features of a typical 
water form. A similar zonation was noticed by Johnson (1986: 26, 41) in a drying 
pond in northwestern Venezuela, where M. ancylopoda A. Br. grew in the emerged 
outer zone, and M. deflexa A. Br. in the central submerged one. 



236 



FERN GAZETTE: VOLUME 1 3 PART 4 (1 988) 




FIGURE 3. Marsilea berhautii: aquatic species with floating shoots and freely exposed sporocarps. 
27 October 1977, J. Kornas PI. Afr. 6282, KRA. 



MARSILEA IN NIGERIA 



237 




FIGURE 4. Marsilea distorta: terrestrial species with geocarpic sporocarps. Plants appearing earlier 
in the dry season (A) are rather mesomorphic, while those appearing later (B) have more 
xeromorphic features. A - 26 October 1977, J. Kornas PI. Afr. 6271, KRA: B - 2 December 
1977, J. Kornas PI. Afr. 6433, KRA. 



The position of sporocarps on the plant not only forms an essential taxonomic 
character of the Marsilea species, but it is also of a great biological importance for 
them. In M. berhautii, the only truly aquatic species, the sporocarps are produced 
in long rows high on the petiole and thus fully exposed, without any kind of protection 
(Fig. 3). Two of the amphibious species are basicarpic, with sessile (M. nubica) or 
shortly pedicellate (M. minuta) sporocarps attached to the base of the petiole, just 
above ground level (Figs. 6, 7). The third amphibious species, M. subterranea, is 
geocarpic, with sporocarps buried underground due to the positive geotropic growth 
of pedicels, which penetrate the clay to a depth of 1 cm or more when it is still 
water-logged and soft (Fig. 8). The only terrestrial species under investigations, M. 
distorta, is also geocarpic (Fig. 4). Both basicarpy and especially geocarpy result in 
the location of sporocarps in the immediate vicinity of the parent plant (atelechory), 
and seriously impede their long-distance dispersal (telechory) which occurs in Marsilea 
through the activity of water birds (Johnson 1 986, Malome & Proctor 1 965). As pointed 
out by Stopp (1958), atelechoric dispersal (which never occurs in true aquatics) is 
known mainly in plants of arid and semi-arid zones. It results in the offspring being 
placed in the same ecological niches in which their parents occur, i.e. under suitable 



238 



FERN GAZETTE: VOLUME 13 PART 4 (1988) 




FIGURE 5. Marsilea minuta: sterile aquatic plant with floating leaves. 16 November 1977, J. 
Kornas PI. Afr. 6378, KRA. 



habitat conditions. This is of course of special importance in the extremely patchy 
environment of the arid and semi-arid zones, where small strictly localized pockets 
with favourable life conditions (pools in the case of Marsilea) are surrounded by vast 
expanses of hostile dry land. 

It is tempting to hypothesize that in the evolutionary history of the genus Marsilea 
in Africa there was a shift in the pedicel attachment resulting in the switch from 
an exposed position of sporocarps in the ancestral hydrophilous species to the basicarpic 



MARSILEA IN NIGERIA 



239 




FIGURE 6. Marsilea minuta: fertile plants with basicarpic sporocarps. A - transitional stage 
between aquatic and terrestrial forms with both floating and aerial leaves: B, C - terrestrial 
plants changing from mesomorphic to xeromorphic features as the drought increases. A - 16 
November 1977, J. Kornas PI. Afr. 6378, KRA; B - 24 November 1977, J. Kornas PI. Afr. 6418, 
KRA; C - 3 February 1978, J. Kornas PI. Afr. 6626, KRA. 



240 



FERN GAZETTE: VOLUME 1 3 PART 4 (1 988) 




FIGURE 7. Marsilea nubica: amphibious species with basicarpic sporocarps. A - sterile, aquatic 
plant with both submerged and floating leaves; B - fertile terrestrial plant. A - 30 September 
1 977, J. Kornas PI. Afr. 6161. KRA; B - 1 6 November 1 977, J. Kornas PI. Afr. 6379, KRA. 




I £r>H?f >\7 






FIGURE 8. Marsilea subterranea: amphibious species with geocarpic sporocarps. 19 January 
1 978, J. Kornas PI. Afr. 6581 , KRA. 



and geocarpic position in the derived xerophytic species, with simultaneous restriction 
of the long-distance (telechoric) dispersal in favour of the atelechoric one. The 
conclusions of Bhardwaja's (1967, 1980) and Gopal's (1968) studies on the Indian 
species of Marsilea are quite consistent with this hypothesis. It fits very well in a 
more general hypothesis of Johnson (1986: 14) that in Marsilea "adaptations toward 
passing a greater portion of the life on land are secondarily acquired, rather than 
primitive, being superposed on a plant form and life cycle designed for life in water". 



MARSILEA IN NIGERIA 



241 




FIGURE 9. Zonation of Marsilea species in a roadside ditch near Gajibo on Maiduguri - Gambaru 
road. A - pool filled with water at the end of the rainy season; B - dried out pool late in the 
rainless season (3 February 1979): a - M. nubica (leafless and completely dry), b. - M. minuta 
(with small living leaves). 



AN OUTLOOK 
There are possibly many more biological features of the Marsilea species in the arid 
and semi-arid zones which contribute to the success of these plants in their harsh 
environment, and certainly deserve a closer study. One of such peculiarities is the 
notorious longevity of sporocarps, which are reported to be able to germinate after 
more than 60 or even 1 00 years (Allsopp 1 952, Bhardwaja 1 980, Bloom 1 955, Johnson 
1 985, Stopp 1 958), and their extreme resistance against environmental stress (drought, 
heat, etc. - Bloom 1961). Another is the hygrochasic mechanism of sporocarp 
dehiscence, connected with the ability of the gametophyte of an immediate fertilization 
and a rapid development of young embryos in the aquatic medium (Johnson 1986: 
25). Still another peculiar feature is the production of drought-resistant vegetative 
propagules ("tubers" consisting of condensed rhizome branches with many small 
leaves), reported to occur not only in the Australian species M. hirsuta R. Br. (Braun 
1873, Goebel 1918, Clifford & Constantine 1980), but also in M. minuta (Bhardwaja 
1980, Gopal 1966, Loyal & Kumar 1979), though not confirmed in the present study. 
Bizarre growth forms in some African species, e.g. in M. botryocarpa (Launert 1968, 
1984) or M. megalomanica (Launert 1970, 1984) suggest that still more unknown 
life-phenomena may be discovered in this fascinating and biologically little explored 
genus. 

ACKNOWLEDGEMENTS 
I am very thankful to my wife, Dr Anna Medwecka-Kornas, and my colleagues from 
the Biology Department, University of Maiduguri, Dr Ivonne Garrod, Mike Pearson 
and Ros Hancock, for their assistance during the collecting trips. I also thank Dr 
E. Launert and Mr A.C. Jermy, both of British Museum (NH), London, for the critical 
revision of voucher material of Marsilea and Isoetes, respectively, as well as Malgorzata 
Matyjaszkiewicz, M.Sc, for drawing the figures. 



242 FERN GAZETTE: VOLUME 1 3 PART 4 (1 988) 



REFERENCES 

ALLSOPP, A. 1952. Longevity of Marsilea sporocarps. Nature (London) 169: 79-80. 

ALLSOPP, A. 1954. Experimental and analytical studies of pteridophytes. XXIV. Investigations 

on Marsilea. 4. Anatomical effects of changes in sugar concentration. Ann. Bot. (London) 

7 g; 449-461. 
ALLSOPP, A. 1963. Morphogenesis in Marsilea. J. Linn. Soc, Bot. 58 (373): 417-427. 
BHARDWAJA, T.N. 1967. Pedicel attachement in Marsilea diffusa var. approximata A.Br, in 

relation to habitat factor. Trop. Ecol. 8: 1 7-21 . 
BHARDWAJA, T.N. 1 980. Recent advances in our knowledge of the water fern Marsilea. Aspects 

of Plant Sciences 3: 39-62. 
BLOOM, W.W. 1955. Comparative viability of sporocarps of Marsilea quadrifolia in relation to 

age. Trans. Illinois State Acad. Sci. 47: 72-76. 
BLOOM, W.W. 1961. Heat resistance of sporocarps of Marsilea quadrifolia. Amer. Fern J. 51 

(2): 95-97. 
BRAUN, A. 1873. Nachtragliche Mitteilungen uber die Gattungen Marsilia und Pilularia. 

Monatsber. Kdnigl. Preuss. Akad. Wiss. Berlin 1872: 635-679. 
CLIFFORD, H.T. & CONSTANTINE, J. 1 980. Ferns, fern allies and conifers of Australia. A laboratory 

manual. XVI + 150 pp. Univ. of Queensland Press, St. Lucia, Queensland. 
COOK, C.D.K., GUT, B.J., RIX, E.M., SCHNELLER, J. & SEITZ, M. 1974. Water plants of the 

world. A manual for identification of the genera of fresh water macrophytes. VIII + 561 

pp. Dr. W. Junk, The Hague. 
GAUDET, J.J. 1 964 Morphology of Marsilea vestita. II. Morphology of the adult land and submerged 

leaves. Amer. J. Bot. 51 (6): 591 -597. 
GLUCK, H. 1922. Uber die knollchenartigen Niederblatter an dem Rhizom von Marsilea hirsuta 

A.Br, und ihre Beziehungen zu den Primar- und Folgeblattern. Flora 1 15: 251 -258. 
GLUCK, H. 1936. Pteridophyten und Phanerogamen. In: Pascher, A. (ed.) Die Siisswasserflora 

Mitteleuropas. Gustav Fischer, Jena, H. 1 5, pp. XX + 486. 
GOEBEL, K. 1918. Organographie der Pflanzen insbesondere der Archegoniaten und 

Samenpflanzen. Zweiter Teil: Spezielle Organographie. Heft 2: Pteridophyten. Zweite 

Auflage. Gustav Fischer, Jena, XI-XII, 903-1208 pp. 
GOPAL, B. 1966. Vegetative propagation in Marsilea minuta Linn. J. Indian Bot. Soc. 45: 352- 

353. 
GOPAL, B. 1968. Ecological studies of the genus Marsilea. I. Water relations. Trop. Ecol. 9: 

153-170. 
GOPAL, B. 1969. Ecological studies on the genus Marsilea. II. Edaphic factors. Trop. Ecol. 10: 

278-291. 
GROVE, AT. 1970. Africa south of the Sahara. XIV + 385 pp. Oxford Univ. Press, Oxford. 
GUPTA, K.M. 1962. Marsilea. Botanical Monograph No. 2. X + 113 pp. Council Sci. Ind. Res., 

New Delhi. 
HUTCHINSON, L.L.D. & DALZIEL, J.M. 1954-1972. Flora of West Tropical Africa. Vol. I: VIII 

+ 828 pp; Vol. II: XI & 544 pp; Vol. Ill: VI + 574 pp. Crown Agents for Overseas Governments 

and Administrations, London. 
JOHNSON, DM. 1985. New records for longevity of Marsilea sporocarps. Amer. Fern J. 75: 

30-31. 
JOHNSON, D.M. 1986. Systematics of the New World species of Marsilea (Marsileaceae). Syst. 

Bot. Monogr. 11: 1-87. 
KORNAS, J. 1983. Pteridophyta collected in Northern Nigeria and Northern Cameroon. Acta 

Soc. Bot. Poloniae 52: 321 -335. 
KORNAS, J. 1985. Adaptive strategies of African pteridophytes to extreme environments. Proc. 

Roy. Soc. Edinburgh 86 B? 391 -396. 
LAND SYSTEMS. 1970. Land systems. North East Nigeria 1: 500 000. Map 6a, 6b. Land Resource 

Study, The Land Resources of North East Nigeria. Land Resource Division of Overseas 

Surveys, London. 
LAUNERT, E. 1968. A monographic survey of the genus Marsilea Linnaeus. I. The species of 

Africa and Madagascar. Senckenberg. Biol. 49 (3/4): 273-31 5. 
LAUNERT, E. 1970. Marsilea Linnaeus: Variation on a theme. Senckenberg. Biol. 51 (5/6): 433- 

435. 
LAUNERT, E. 1971. A remarkable new African species of Marsilea Linnaeus. Seckenberg. Biol. 

52 (6). 449-452. 
LAUNERT, E. 1984. A revised key to and new records of African species of the genus Marsilea. 

Garcia de Orta, Ser. Bot. 6 (1-2): 1 1 9-140. 



MARSILEA IN NIGERIA 243 



LOYAL, D.S. & KUMAR, K. 1 979. Cytological observations on some natural populations of Marsilea 

minuta L. - their reproductive mechanisms and evolutionary future. In: Bir, S.S. (ed.) 

Recent Researches in Plant Sciences. Kalyani Publishers, New Delhi, pp. 229-236. 
MALONE, C.R. & PROCTOR, V.W. 1965. Dispersal of Marsilea mucronata by water birds. Amer. 

Fern J. 55 (4): 1 67 -170. 
SADEBECK, R. 1902. Marsiliaceae. In Engler, A. & Prantl, K. (eds.) Die naturlichen 

Pflanzenfamilien, Teil. I, Abt. Ill, Embryophyta Siphonogama, Unterabteilung 2. Pteridophyta. 

W. Engelmann, Leipzig, pp. 403-421. 
SCHELPE, E.A.C.L.E. & ANTHONY, N.C. 1986. Flora of Southern Africa. Pteridophyta. XV + 292 

pp. Bot. Res. Inst., Dept. Agric. Water Supply, Republic of South Africa, Pretoria. 
SCHMIDT, K.D. 1978. Ein Beitrag zum Verstandis der Morphologie und Anatomie der 

Marsileaceae. Beitr. Biol. Pflanzen 54 ( 1): 41 -91. 
STOPP, K. 1958. Die verbreitungshemmenden Einrichtungen in der sudafrikanischen Flora. Bot. 

Stud. 8. 1-103. 
TRYON, R.M. & TRYON, A.F. 1982. Ferns and allied plants with special reference to tropical 

America. XII + 857 pp. Springer Verlag, Berlin. 
WALTER, H. 1971. Ecology of tropical and subtropical vegetation. XVIII + 539 pp. Oliver & Boyd, 

Edinburgh. 
WALTER, H. & LIETH, H. 1960, Klimadiagramm-Weltatlas. G. Fischer, Jena. 
WHITE, F. 1983. The vegetation of Africa. Vegetation map of Africa. 356 pp. UNESCO, Paris. 
WHITE, R.A. 1966. The morphogenetic effects of protein synthesis inhibition in Marsilea. Amer. 

J. Bot. 53:158-165. 
WILLIS, J.C. 1973. A dictionary of the flowering plants and ferns. 8th ed., rev. by H.K. Airy 

Shaw. XXII + 1245 + LXVI pp. Cambridge Univ. Press, Cambridge. 



244 FERN GAZETTE: VOLUME 1 3 PART 4 (1 988) 



FERN GAZ. 13(4)1988 245 



SHORT NOTES 



DRYOPTERIS VILLARH (DRYOPTERIDACEAE: PTERIDOPHYTA) A NEW 
HIGH-MOUNTAIN SPECIES IN THE CARPATHIANS 



In 1 986, when collecting field data for the "Atlas of distribution of the vascular plants 
in the Tatra National Park" (Western Carpathians, Poland) we found one plant of 
a fern unknown to us. As this was in very late October, after the first ground frosts 
in the mountains, the plant was badly damaged and with no spores on it. In this 
state certain identification was not possible. Next year, 1987, very careful field 
searching in the same region resulted in finding three more specimens, this time 
in very good condition. Their gross morphology (Fig. 1), the dense indumentum of 
glandular hairs on both sides of fronds and some other features indicate that they 
belong to Dryopteris villarii (Bellardi) Woynar ex Schinz & Thell. subsp. villarii. 

The locality is in the Polish part of the Tatra Mts, in the postglacial hanging valley 
called "Swistowka Wielka" above the "Wantule" strict reserve (upper part of the 
Dolina Mietusia valley), at an altitude of 1360m, in the Pinus mugo (subalpine) belt 
(considerably lowered in this region because of orographic conditions). The species 
occurs there on limestone block scree of fairly large boulder size, in an open situation, 
on a slightly (5°) inclined northern slope. The rather moist soil, consisting of mineral 
grains intimately mixed with well-decomposed black humus, develops there in local 
pockets among boulders. The very small population of D. villarii (only three plants 
on about one square metre) was accompanied by the following species: Aconitum 
variegatum, Allium montanum, Asplenium viride, Bellidiastrum michelii, Carduus 
glaucus, Carex sempervirens, Delphinium oxysepalum, Galium anisophyllum, Poa 
alpina, Rhodiola rosea, Saussurea alpina, Saxifraga aizoides, S. moschata, S. 
paniculata, Scabiosa lucida, Selaginella selaginoides, Solidago alpestris, Thalictrum 
minus, and Viola bi flora. 

The locality of D. villarii in the Tatra Mts is the first site not only for Poland and 
for this part of Europe, but also for the Carpathian Mts as a whole. The nearest 
known stands of this species in the Eastern Alps (Fraser-Jenkins 1 977, Fraser-Jenkins 
& Reichstein 1984) are about 500km distant from the locality discovered. 

The authors are greatly indebted to Dr W.H. Paryski for improving the English 
style of the manuscript. 

REFERENCES 
FRASER-JENKINS, C.R. (1977). Three species in the Dryopteris villarii aggregate. Candollea 32: 

305-319 
FRASER-JENKINS, C.R. & REICHSTEIN, T. (1984). Genus Dryopteris Ada ns. In: K. U. Kramer 

(ed.), Pteridophyta in G. Hegi, lllustrierte Flora von Mitteleuropa, 1(1), (3rd ed.). Berlin 

& Hamburg. 

HALINA PIEKOS-MIRKOWA 

Nature Protection Research Centre, Polish Academy of Sciences, 

Smolensk 14,31-112 Krakow, Poland. 

ZBIGNIEWMIREK 

Institute of Botany, Polish Academy of Sciences, 

Lubicz 46, 31 -51 2 Krakow, Poland. 



246 



FERN GAZETTE: VOLUME 1 3 PART 4 (1 988) 




3bMtf£ 




Fig. 1 . Fronds of Dryopteris villarii subsp. villarii from the locality in the Tatras. 



LYGODIUM JAPONICUM IN SINGAPORE 



Holttum (Flora Malesiana II, 1(1), 1959) says Lygodium japonicum (Thunb.) Sw. is 
native only to regions with a pronounced dry season, and Alston & Holttum (Reinwardtia 
5, 1959) reported that it will grow in cultivation in Singapore but not vigorously, 
and that in its natural habitat the fronds probably die out during the dry season, 
to appear again with the beginning of the wet period. 

For the last five years or more L japonicum has apparently been growing profusely 
around the periphery of the lawn as well as along the fence fronting a house occupied 



FERNGAZ. 13(4)1988 247 



for years by expatriate Japanese at Chun Tin Road, Singapore. These plants are 
evergreen and fertile most of the time, encouraged presumably by the wet climate. 

We are grateful to Professor R.E. Holttum of The Royal Botanic Gardens, Kew 
for confirming the identification of our specimen. 

Y.C. WEE & LL CHUA 
Dept of Botany, National University of Singapore, Singapore 051 1, 

Republic of Singapore. 

DRYOPTERIS X GOMERICA 
(DRYOPTERIDACEAE: PTERIDOPHYTA), NEW FOR EUROPE 

THE DISCOVERY 

In September 1 986 the author, A.C. Jermy (BM) and M. Hermy (INC, Hasselt, Belgium) 
met Prof. M. Mayor-Lopez and H. Nava (Oviedo) for a reconnaissance trip in Asturias, 
N. Spain, to prepare an excursion-guide for the XlVth International Botanical Congress 
in Berlin (Viane, Mayor-Lopez & Jermy 1987). Asturias was chosen since it is 
particularly rich in pteridophytes (c. 66 taxa), with arctic-middle european (c. 30%), 
atlantic (c. 26%), mediterranean (c. 23%), macaronesian (c. 13%) and endemic 
(c. 5%) taxa (see also Fraser-Jenkins & Gibby 1986). The wet and mild climate on 
the Mirador del Fito (600m alt.) in the Sueve Mts., north of Arriondas and only c.4km 
from the Gulf of Biscay supports a mixture of most interesting ferns, including Culcita 
macrocarpa C. Presl (Fraser-Jenkins & Lainz 1 983). The acid, greyish-white sandstone 
is covered with Ulexgallii - Erica arborea - E. mackaiana heathland and some isolated 
groups of Betula, Pinus (planted) and Eucalyptus (planted). This site is probably the 
only locality in northern Spain where both Dryopteris aemula (Ait.) 0. Kuntze and 
D. guanchica Gibby & Jermy grow together abundantly. It was here, at the base 
of some large boulders and well protected against grazing by an almost impenetrable 
spiny mass of Ulex, that a large plant, intermediate in morphology between D. aemula 
and D. guanchica, was collected (R. Viane 3355). Part of the rhizome was taken 
for cultivation and subsequent cytological study. Closer observations, showing that 
the plant has aborted spores and that micromorphological characters are also 
intermediate between D. aemula and D. guanchica, led us to identify the plant as 
D. x gomerica. Fixations for cytological analysis were taken in May 1987 and sent 
to H. Rasbach (Glottertal, W. Germany) who reported (in litt. 27.9.1 987 & 1 1 .1 1 .1 987) 
that the plant is triploid with n = c. 41" and 41 ! at meiosis. This result not only 
confirms our identification but is also in agreement with the results of Gibby & Widen 
(1983). 

Hitherto D. x gomerica was only known from La Gomera (Canary Isl.), apparently 
the only other locality where both parents grow together. Unlike the Azores, the Canary 
Islands (and Madeira) are not included in the area covered by the Flora Europaea, 
thus our new Asturian locality is the first record for mainland Europe. 

THE DISTINCTION 

Cytological, chemical and morphological evidence have led to the present interpretation 
of the relationships between D. x gomerica and its parents (Gibby et al. 1978; Gibby 
1979; Gibby 1983; Gibby & Widen 1983). D. guanchica is an allotetraploid species 
that originated via chromosome doubling of a hybrid between D. intermedia ssp. 
maderensis and D. aemula. D. x gomerica thus represents a "backcross" of D. guanchica 
with its D. aemula ancestor (fig. 1); consequently the overall morphological distinction 



248 



FERN GAZETTE: VOLUME 13 PART 4 (1988) 



between these taxa is fading out through the morphologically intermediate hybrid 

(fig. D- 

Some characters for identification are given in the table below. All values are 
in Aim; they represent the mean ± the standard deviation; "glands" means the 
unicellular capitate hairs, "hair" means all uniseriate multicellular hairs. 

Abbreviations used (hairs): Lt = total length; Lc = length apical cells; Wc = width 
apical cells; N = number of cells in a hair. For the glands: It =3 total length; dA 
= diameter of apex; dB = diameter of base; Is = stalk length. 



MACROSCOPIC CHARACTERS 



D. aemula D. x gomerica 

plant habit fronds: flat, drooping fronds: erect 

frond colour medium to yellow-green yellow-green 



pinna flat with drooping tip 

pinnules concave: edges and 

whole ultimate segment 
turned up 

pinna apex attenuate to caudate 



flat 

flat with only the edge 
of the segment and its 
ultimate tip turned up 

caudate 



D. guanchica 

fronds: erect 

blue(metallic) to dark green 
(N.Spain) 

flat (occas. convex when 
pinnules reflexed) 

flat, rarely slightly convex, but 
with the extreme segment tip 
turned up 

caudate 



MICROSCOPIC CHARACTERS 





D. aemula 


D. x gomerica 


D. guanchica 


cytology 


diploid: n = 41" 


triploid: n = 41" &41 1 


allotetraploid: n = 82" 


spores 


good 


aborted clumps 


good 


exospore 


34 ± 2/um long 




34 ± 2;um long 


perispore 


costate-venate 


irregular 


(narrowly) costate-echinulate 


sori 


with paraphyses = 
laminal hairs 


no paraphyses 


no paraphyses 


sporangia 


2-cellular gland-tipped 


glabrous, rarely with 


glabrous, rarely with a gland 




hair or unicellular gland 


a gland 






at the base 






indusium 








dorsal 


glandular or glabrous 


glands 


glands 


marginal 


glands and (often) hairs 


glands and hairs 


glands only 


stomata 


39-44-49/um long 


45-48-52/um long 
lamina indument 


44-49-54>um long 


small paleae 


slightly bullate with 
± isodiametric cells and 
a long uniseriate apex 


intermediate 


not bullate, cells elongate 


hairs: 








Lt 


210-360-510 


450 - 660 - 750 


250 - 450 - 700 


Lc 


30-50 


37-57 


36-54 


Wc 


18-28 


34-36 


31 -45 


Lt/Wc 


1.3-2.1 


1.5-1.9 


1.1 - 1.5 


N 


7-13 


9- 15 


8- 14 


glands: 








It 


53 ±7 


73 ±6 


76 ±8 


dA 


43 ±5 


48 ±3 


38 ±5 


dB 


15±2 


17± 1 


18±2 


Is 


17±5 


25 ±6 


36 ±8 


dA/ls 


2.7 


1.9 


1.1 


dA/dB 


2.9 


2.8 


2.1 



FERNGAZ. 13(4)1988 



249 



\ 






D.aemula (AA) 




D guanchica(A Al I) 



D.xgomerica(AAI 



FIG. 1 Silhouettes of Dryopteris x gomerica and its ancestors. 

REFERENCES 
FRASER-JENKINS, C. R. & LAINZ, M. 1983. Culcita macrocarpa - a new locality in Spain. Fern 

Gazette 12: 299-300. 
FRASER-JENKINS, C. R. & GIBBY, M. 1986. A new Dryopteris hybrid from Spain. Fern Gazette 

73/113-116 
GIBBY, M. 1979, Palaeoendemism and Evolution in Macaronesian Dryopteris. In. D. Bramwell 

(Ed.), Plants and Islands. Academic Press, London-New York. 
GIBBY, M. 1983. The Dryopteris dilatata complex in Macaronesia and the Iberian Peninsula. 

Acta Bot. Malac. 8: 59-72. 
GIBBY, M. & WIDEN, C.-J. 1983. Three new Dryopteris hybrids from Spain and the Canary 

Islands. Fern Gazette 12: 267-270. 
GIBBY, M., WIDEN. C.-J. & WIDEN, H. K. 1978. Cytogenetic and phytochemical investigations 

in hybrids of macaronesian Dryopteris (Pteridophyta: Aspidiaceae). Plant Syst. & Evol. 

730.235-252. 
VIANE, R., MAYOR -LOPEZ, M. & JERMY, C. 1987. Excursion 39. The systematics and ecology 

of the Pteridophytes of northern Spain. Excursion guide: XlVth International Botanical 

Congress, Berlin. 

R. VIANE 

Laboratorium voor Morfologie, Systematiek en Ecologie der Planten 



K. L Ledeganckstraat 35, 9000 GENT, Belgium. 



250 FERN GAZETTE: VOLUME 1 3 PART 4 (1 988) 



REVIEWS 

LIEBM 'ANN'S MEXICAN FERNS by J.T. Mickel, R. McVaugh, S. Karell and H. Balslev. 
350 pp. Contributions from the New York Botanical Garden volume 19. 1987. ISBN 
89327 324 4. Price $30.50 for U.S. orders, $31.75 for non-U.S. orders. 

Liebmann's Mexican Ferns consists of an Introduction by John Mickel, an Itinerary 
and Gazetteer by Rogers McVaugh, a translation of Liebmann's "Mexicos Bregner" 
by Sven Karell and Henrik Balslev, an index to the translation, a reprinting of Liebmann's 
"Mexicos Bregner" (in Danish) and an index to "Mexicos Bregner". Liebmann's 
"Mexicos Bregner" has for a long time been inaccessible to the average botanist 
both on account of its rarity as a publication - it is only held in major botanical 
libraries - and because although the plant descriptions are in Latin the important 
field observations and comments are in Danish. It is a work of great importance 
to students of Mexican ferns, being the second of the four Mexican fern floras produced 
last century. Like the first of these floras (Martens & Galeotti, Memoire sur les Fougeres 
du Mexique, 1 842), Liebmann's work (1 849) was based on field experience in Mexico, 
unlike the two later works (Fee, Catalogue methodique des fougeres et des 
Lycopodiacees du Mexique, 1857 and Fournier, Mexicanas plantas. Vol. 1. 
Cryptogamia, 1872) which were prepared in Paris and based only on herbarium 
collections. Liebmann provided information on 308 species, 95 of which were described 
as new. 

The authors are to be congratulated upon making Liebmann's work more widely 
available both by translating it and reprinting the original. The addition of modern 
names and synonymy to the translation makes it usable with the modern fern floras 
available for some Mexican states while the itinerary and gazetteer will be of great 
use to students of Liebmann's collections, whether of ferns or higher plants. 

B.S. PARRIS 



FERN GAZ. 13(4)1988 251 



FERNS OF MALAYSIA IN COLOUR by AG. Piggott. 458 pp. Tropical Press, Malaysia. 
ISBN 967 73 0029 6. 1988. Price £43.00, U.S. $69.00 including surface mail postage 
or M$ 150.00. 

This beautiful book is the product of many years work by Audrey Piggott and her 
husband John who took the photographs. It is intended as a companion volume to 
Flora of Malaya Volume II Ferns of Malaya by Dr RE. Holttum (1954 and 2nd edition 
1968), and, in spite of its rather misleading title which led to a colleague purchasing 
the book under the misapprehension that he would be able to identify East Malaysian 
(Sabah and Sarawak) ferns, its coverage is effectively that of Peninsular Malaysia 
and Singapore, i.e. that of Holttum's original work. The addition of Phanerosorus, 
a genus which occurs in East Malaysia but not in the Peninsula, I find irrelevant 
as numerous other fern genera with a similar distribution could well have been included 
but would have lessened the book's value as a companion to Ferns of Malaya. In 
fact many of the species illustrated here also occur in Sabah and Sarawak although 
their occurrence in either or both of these states is only mentioned for six taxa. 

The format consists of an introduction, a summary of the life cycle of a fern, 
the principal vegetation types and fern habitats, pests and diseases of ferns and the 
main account of the ferns which are arranged in a taxonomic order based partly 
on Holttum, partly on Flora Malesiana revisions and partly on Pichi Sermolli's generic 
arrangement. With two exceptions, Monachosorum and Polystichum, all fern genera 
currently known to occur in Peninsular Malaysia are included in Piggott's book. Of 
the 598 fern taxa in the area (Parris, unpublished data) 392, or c. 66%, are illustrated. 
The discrepancy between this figure and the nearly 80% coverage claimed by Mrs 
Piggott is largely due to numerous very recent revisions and additions to the flora 
which have appeared since the book went to press, judging by the date of April 1984 
at the end of the acknowledgements. These include Bidin, Fern Gazette 12: 360- 
361, 1984 (Osmunda vachellii), Croxall, Kew Bull. 41:519-531, 1986 (new records 
of Microgonium (= Trichomanes p.p. of Piggott and Holttum), Holttum, Gardens' Bull. 
38:145-148, 1985 (two new species of Tectaria) and Parris, Kew Bull. 41:491-517, 
1986 (new species and new records of Grammitidaceae). Surprisingly one species 
described in this last paper has found its way into the book, however. 

The name used in Holttum and the page number where the species is described 
are given for easy reference. Where the currently accepted name is different from 
that in Holttum, the name he used is also given, although no distinction is made 
between taxonomic synonyms and misidentif ications by Holttum which should correctly 
be cited as 'sensu Holttum' rather than being attributed to their authors. The 
nomenclature is on the whole up to date with that used in revisions available at 
the time of writing although, inexplicably, three names in Lindsaea have not been 
brought up to date with those used by Kramer in his account of the genus for Flora 
Malesiana although others have been corrected. Lindsaea scandens should be L. 
parasitica. L nitida should be L Integra and L decomposita should be L. cultrata. 
Other incorrect names used are Trichomanes proliferum for T. minutum, Asplenium 
squamulatum for A. vittaeforme, A. adiantoides for A. polyodon, Tectaria oligophylla 
for T. fissa and Blechnum indicum for B. serrulatum. 

For almoLt every taxon included there is a habitat photograph, a photograph of 
a frond and a close-up photograph showing details of the frond which usually includes 
the sori, together with distribution notes, a short description and medicinal uses where 
known. The photographs are of good quality on the whole and show various diagnostic 
features. The inclusion of traditional medicinal uses adds to the interest of the book. 
Many scientists are now concerned with recording such information and testing the 



252 FERN GAZETTE: VOLUME 1 3 PART 4 (1 988) 



efficacy of 'bush medicines' and this compilation will doubtless be a useful aid to 
ethnobotanists working in the peninsula. 

Our state of knowledge of the fern flora of the Malaysian Peninsula is far from 
complete, as shown by the number of taxa discovered in or described from the area 
since this book was written. I am preparing descriptions of another three new species 
from the peninsula (one Coryphopteris and two Grammitis); the first was collected 
by me during field work in 1986 and is not represented otherwise by herbarium 
specimens while the other two have been known and wrongly identified in herbaria 
for several years. There are probably many more such additions to be made to the 
flora both by field work and by detailed studies on existing herbarium specimens, 
but they will not detract essentially from the usefulness of Holttum's classic work 
on Malayan ferns, which remains one of the best fern floras ever written, or from 
Piggott's book which with its colour photographs and nomenclature updated to c. 
1983 will be a good companion volume. I would recommend this book, in fact, to 
everyone who has a 'Holttum' and an interest in ferns in this part of the world. 

B.S. PARRIS 



FERN GAZ. 13(4)1988 253 

OBITUARY 

IRENE MANTON (1 904-1 988) 

Emeritus Professor Irene Manton, F.R.S., for thirty years an honorary member, and 
past-President (1969-1972), of the British Pteridological Society, died in hospital in 
Leeds on 31 May 1 988, after a brief illness. 

The younger of two eminent biologist daughters of a London dentist, Irene Manton 
was educated at St. Paul's Girls' School, and at Girton College, Cambridge, where 
she undertook both undergraduate (1923-6) and postgraduate (1926-9) studies. She 
had become fascinated with chromosomes whilst still at school, and the opportunity 
to indulge her interest came when she spent the first year of her postgraduate research 
in Stockholm in 1926-27. Her early work was on the cytology of the Cruciferae and 
she came to a study of ferns almost accidentally, through a happy combination of 
her passion for chromosomes and her appointment in 1 929 to an assistant lectureship 
in the Botany Department of Manchester University where the then Professor of 
Cryptogamic Botany, W.H. Lang (himself trained by the famous pteridologist, F.O. 
Bower, and discoverer with Kidston of the early fossil pteridophyte genus, Rhynia), 
was using Osmunda in experimental studies on the effect of induced apogamy on 
the cytological basis of alternation of generations. She studied the cytology of Lang's 
apogamously produced plants and in the process discovered not only another polyploid 
series (she had found her first in the watercress) but also the delights and technical 
tractability of the Osmundaceae as cytological subjects. This led to her series of 
important papers on the spiral structure of the chromosomes in the Royal Fern, 
Osmunda regalis. Other ferns, however, were not so easy, and proved to be such 
difficult cytological material that she immediately determined to conquer them. During 
the war years she worked tirelessly on the British pteridophyte flora and, when the 
war came to an end, she had accumulated so much data that she decided to publish 
her results in book form, Problems of Cytology and Evolution in the Pteridophyta 
(1950). She had by this time moved to Leeds, having been appointed to the Chair 
of Botany in January 1946, and it was here that she took up her next challenge. 
The recently invented electron microscope, by the great increase in resolution which 
it provided, had opened up boundless possibilities for the microscopist, and Manton 
responded by setting up the first, which was for some time the only, laboratory in 
the world to study the ultrastructure of plants. Electron microscopy, with which she 
was to be principally involved for the whole of the rest of her life, became her new 
consuming passion, while the fern work was continued by her colleagues, research 
students and visiting postdoctoral research fellows. She was an exacting, and at times 
intimidating, person to work for, as she had high standards and did not tolerate fools 
glady: on occasions we all felt the rough edge of her tongue. But we knew also 
the other side: her warm, affectionate nature, her extreme kindness and generosity 
to those of her 'family' in need and her well-developed sense of fun. She had a 
vigorous intellectual curiosity, and throughout her life she retained a child's infectious 
delight in the thrill of making discoveries. She had a quick, incisive mind, great energy, 
an enormous capacity for hard work as well as boundless enthusiasm. Her research 
was her life, and she cared little for such things as food, clothes and holidays. Botany 
House during the 1960's was an exciting place to be. As professor she was busy 
during the working day with her teaching duties and with administration, but every 
evening and weekend she worked on her electron microscope. There was a wonderful 
atmosphere of active research that permeated the department, and her colleagues 
and students were stimulated to work similar ungodly hours. Indeed, we felt guilty 
about taking an hour off on Saturday afternoons to buy food for Sunday's lunch! 



254 FERN GAZETTE: VOLUME 13 PART 4 (1988) 



However, it was not only botanical research to which she devoted her abilities. 
Sometimes, during the evening, she would take a break for coffee, and such occasions 
were an opportunity for her to show us, with characteristic delight and pride, her 
latest acquisition, whether it was a fragment of papyrus text, a piece of Roman 
glassware or a new painting. Her life-long interest in form and shape led her far 
beyond the bounds of science and into the world of art. She was an avid collector 
of artifacts, prints and paintings which she bought as they took her interest, and 
not at all for their intrinsic value. These were used to illustrate lectures and talks 
and especially to, stimulate the young, (her own expression). A naturally gifted person, 
who, in addition to her other skills, was a talented violinist, she appears, from a 
very early age, to have had a keen sense of history, particularly as it applies to biology 
(an early article, written whilst still a student, describes the history of the vine, and 
is illustrated by motifs from Greek pottery), and she was able to weave art and science 
together in a novel and entertaining manner. She also had a highly original way 
of viewing life and one of her most stimulating series of lectures was entitled 'Other 
Ways of Looking at Nature', which was illustrated by reproductions of cave paintings 
as well as by modern abstract and by Chinese art. A natural story-teller, she held 
her lecture-audiences spell-bound from beginning to end. Her innumerable scientific 
papers are written in the same inimitable, racy style. In 1969 she retired from the 
Chair of Botany at the University of Leeds and, with characteristic determination and 
energy, embarked on what was to be another nineteen years of active and fruitful 
research. 'Life begins at 65' well describes her retirement, which included major 
expeditions to Greenland and the Galapagos Islands in order to collect her beloved 
nanoplankton. 

Irene Manton had a long association with the British Pteridological Society which 
she joined in 1936. In the summer of 1938 she went on their Annual excursion 
to Kenda. Having already discovered different chromosome numbers in the fern Lastrea 
(= Dryopteris) filix-mas, she was hoping to find wild material of 'var. abbreviata (DC) 
Newman' so that she could count its chromosomes, and participants at the meeting 
were able to advise her where she could find it. It is interesting to note that generations 
of Manton's research students, raw at the outset of their postgraduate studies, have 
since followed the same path, similarly having been guided to either plants or localities 
by the kindness and field experience of Society members. Manton's early investigations 
on hybrid Dryopteris in Britain are published in the Gazette of December 1938, and 
the continuation of this work and her extension of it to the rest of the British fern 
flora, culminating in the publication of her book in 1 950, have already been mentioned. 
Preliminary data from the island of Maderia, also included in the book, laid the 
foundation for the comparative cytological analysis of fern floras in different parts 
of the world while the extension of the investigation to the tropics began in late 
1950 with the Leeds University Botany Department's expedition to Ceylon, where 
some 70% of its fern flora was cytologically sampled during the course of this one 
visit; as in the temperate floras already examined, evidence of evolution through 
polyploidy and hybridization was revealed. In Ceylon too Manton first met Professor 
R. E. Holttum of Singapore and their subsequent friendship and collaboration led 
not only to her study of the chromosomes of Malayan pteridophytes and to the use 
of chromosome numbers in tracing fern phylogeny but also to the successful application 
of cytogenetic methods to various groups of tropical ferns. Finally, her pioneering 
work utilising the pairing behaviour of the chromosomes in hybrids to unravel the 
evolution of and the relationship between species in polyploid complexes has been 
taken up and used extensively in the elucidation of difficult groups in the fern floras 
of Europe, North America and Australasia. Manton's own lively description of how 






FERN GAZ. 13(4)1988 255 



her involvement with ferns came about (her Presidential Address to the Society in 
October, 1972) is published in two parts in Gazettes 70(6), 1973 and 10(7), 1974 
while numbers 10(2), 1969 and 10(3), 1970, issued to commemorate her retirement 
from the Chair of Botany at Leeds in 1969, carry articles by her colleagues and former 
research students. Lovis (Gazette 70(1), 1968) details the methods originally devised 
by Manton for the hybridization of ferns, whilst her own last note for the Gazette 
(73(3), 1987) records her friendship and collaboration with Professor T. Reichstein 
of Basel, an association which has resulted in the successful cytogenetic analysis 
of virtually all the European ferns. 

As a scientist, Irene Manton's great strengths were the ability to see the potential 
in a new method or instrument, and the development and adaptation of it to suit 
her own material or interests. A recurring theme throughout her life was her 
exploitation of new techniques. Having in her early cytological research used laborious 
nineteenth-century methods of staining and sectioning, she was quick to realise the 
importance of the acetocarmine squash technique (combined with the McClintock 
method for making slides permanent). Both of these she applied very successfully 
to fern chromosomes, and was able to establish reliable counts, even in high polyploids, 
for the first time. Later on, the use of snail cytase to break down cell-walls enabled 
root-tip squashes to be routinely made and analysed. She also developed photographic 
methods for recording her results and never returned to the drawings with which 
she started at the outset of her career. She was one of the earliest microscopists 
to exploit for biological purposes the increased optical resolution available through 
the use of the ultra-violet microscope, and the enthusiasm with which she took up 
electron microscopy at the earliest possible opportunity is a story too well-known 
to be reiterated here. 

Her remarkable career was recognized by her election to Fellowship of the Royal 
Society (1961) and to the Presidency of the Linnean Society (1973-76) as well as 
to honorary membership of various scientific societies on both sides of the Atlantic. 
She has been awarded honorary doctorates from the universities of McGiil (1958), 
Oslo (1961), Durham (1966), Lancaster (1979) and Leeds (1986) and is the holder 
of numerous prizes and medals for her work. 

It has been a privilege to have known her personally and to have worked under 
her supervision, and her many colleagues and friends will, while they mourn her 
passing with sadness, remember her with great affection. 

ANNE SLEEP 



256 FERN GAZETTE: VOLUME 1 3 PART 4 (1 988) 



THE FERN GAZETTE 

Original papers, articles or notes of any length on any aspect of pteridology will be 
considered for publication. 

Contributions should be sent to: 

The Editor of the Fern Gazette, British Pteridological Society, c/o Department 

of Botany, British Museum (Natural History), Cromwell Road, London SW7 5BD. 

The last date for receiving notes and articles to make the following summer number is: 

31st December each year 

Authors should follow the general style of this number. Close adherence to the following 
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NOTES FOR CONTRIBUTORS 

Manuscripts: Copy should be in English and submitted in double-spaced type with adequate 
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Abstract: All papers, other than short notes, should include a short abstract, to be set at the 
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Headings and sub-headings:These should follow the style of this number. (Primary sub-headings 
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unavoidable for clarity not in the title. All latin names should be underlined throughout the 
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or diagrams in ink, or photographs which must be black and white, and of good technical 
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Photographs should be of the required magnification or larger and need not be made up to 
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British Museum (Natural History), Cromwell Road, London SW7 5BD; for review in 
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Starkes, Ludlow, Shropshire SY8 2HP. 



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The British Pteridological Society 
THE FERN GAZETTE 

VOLUME 13 PART 4 1988 

CONTENTS 

Page 

MAIN ARTICLES 

A chromosome count from Azolla filiculoides (Azollaceae: Pteridophyta) 

- You-Xing Lin & Anne Sleep 1 93 

Cytological and anatomical observations on Tmesipteris 
(Tmesipteridaceae: Pteridophyta) species from New Caledonia 

- A.F. Braithwaite 199 

Shoot temperature measurements of montane Cyathea 
(Cyatheacea: Pteridophyta) species in Papua New Guinea 

- M.J. Earnshaw, T.C. Gunn & JR. Croft 209 

Pteridophytes of Zarate, a forest on the western side of the Peruvian Andes 

- Blanca Leon & Niels Valencia 21 7 

A field survey of Pteridium aquilinum (Dennstaedtiaceae: Pteridophyta) mycorrhizas 

- H.M. Jones & E. Sheffield 225 

Adaptive strategies of Marsilea (Marsileaceae: Pteridophyta) 
in the Lake Chad basin of N.E. Nigeria 

- Jan Kornas 231 



SHORT NOTES 

Dryopteris villarii (Dryopteridaceae: Pteridophyta) a new high-mountain species 

in the Carpathians 

- Halina Piekos-Mirkowa & Zbigniew Mirek 245 

Lygodium japonicum in Singapore 

- Y.C. Wee & L.L. Chua 246 

Dryopteris x gomerica (Dryopteridaceae: Pteridophyta) new for Europe 

- R. Viane 247 

REVIEWS 250 

OBITURY: Irene Manton 

- Anne Sleep 253 

(THE FERN GAZETTE Volume 13 Part 3 was published on 28th July 1988) 



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THE BRITISH PTERIDOLOGICAL SOCIETY 
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FERN GAZ. 13(5) 1989 257 



THE HISTORY OF DIPHASIASTRUM ISSLERI (LYCOPODIACEAE) 
IN BRITAIN AND A REVIEW OF ITS TAXONOMIC STATUS* 

A.C. JERMY 

British Museum (Natural History), Cromwell Road, London SW7 5BD 

ABSTRACT 

The history and distribution in Britain of the plant known today as Diphasiastrum issleri 
is reviewed. In the past there has been much confusion over its identity and relationship 
to D. alpinum and D. complanatum. The diagnostic characters are described and taxonomic 
relationships reconsidered. It is proposed because of apparent introgression between the 
taxa and the likely hybrid origin of D. issleri, that it and D. alpinum be regarded as 
subspecies of D. complanatum and the necessary new nomenclatural combinations are 
made. 

HISTORY 
A taxon distinct from the common alpine clubmoss (Diphasiastrum (Lycopodium) .alpinum) 
was first recognised as a British plant (as Lycopodium complanatum L.) in an editorial 
note in the Gardeners' Chronicle for ll August, 1866 (p. 753) reporting a communication 
from John Lloyd, claiming "it was found by a woman named Sarah Young, while occupied 
in cutting Heath for broom-making", at Lower Waggoner's Wells, Bramshott, Hants. It 
was presumably this record that Charles Cardale Babington (1867) refers to in edition 6 
of his Manual of British Botany, adding "I have not seen it, and doubt its being correctly 
named". However, a year later John Lloyd sent specimens to the editor of the Gardeners' 
Chronicle (Lloyd 1867a) labelled Lycopodium alpinum, from Bramshott, "where the soil 
is a sandy peat, the elevation not more than 600 or 700 feet, the situation sheltered, growing 
near the roots of Heath about 18 inches high, by which it appears to be almost smothered". 
An editorial comment followed this, saying its exact identity is not certain as the specimen 
was sterile and that Professor Babington "inclined to place it under L. chamaecyparissus 
A.Br." (a taxon described from N. America and also found in Europe, very closely related 
to L. complanatum and regarded as synonymous with it at that time). Lloyd commented 
that the main creeping stem was beneath the soil and believed that L. alpinum produced 
stems above the soil. However, in the Chronicle seven weeks later he wrote (Lloyd 1867b) 
that he found L. alpinum with subterranean stems in Wales and therefore believed this 
character not to be important. In edition 7 of the Manual, Babington (1874) still referred 
to the plant as L. complanatum but it is omitted entirely from the 8th edition (Babington 
1881). Later, in a four-page Addendum to the Manual Babington (1883) had obviously 
reconsidered his opinion and cites L. complanatum from Hants., Gloucester, Worcester, 
Ross and Skye. Watson (1870) also recorded the Bramshott plant as L. complanatum but 
also gave "Worcestershire (?), error (?) Bab. Man., ed. 6, p. 445. Leefe msc." According 
to Druce (1882) C.C. Babington wrote to him saying the Rev. Churchill Babington found 
"in July, 1838, on Hartlebury Common, Worcestershire, a Lycopodium which he thought 
might be L. complanatum or chamaecyparissus but of late considered to be only a form 
of L. alpinum, growing at a rather low situation". Babington did not mention a Miss 
Lea (later Mrs Waller), who accompanied him that day, and that she also collected a specimen 
of the Lycopodium (Lees 1867). Her herbarium has not been located (cf. Kent & Allen 
1984), but a sterile scrap of a specimen from Hartlebury is in CGE, however, and appears 
to be D. issleri. 

The next and possibly most significant find was by the Rev. Henry Peter Reader who 
in 1881 collected in Woodchester Park, near Stroud, Gloucs., a plant he thought was L. 
alpinum. A specimen, which had very flattened suberect branches and cones on short 

* This F aper was originally submitted for Pan 3 dedicated to Professor Doctor TADEUS REICHSTEIN 
whose prolific studies on European ferns are a stimulus to many, including the author. 



258 FERN GAZETTE: VOLUME 13 PART 5 (1989) 



peduncles, was sent via Bolton King to Druce, who was stimulated to publish (Druce, 
1882) a note "On Lycopodium complanatum as a British plant" 

The Gloucestershire plant was lent by Druce to Boswell-Syme who had a drawing made 
of it by N.E. Brown for English Botany (Boswell-Syme 1886). The Plate is labelled Lycopodium 
alpinum L. var. decipiens, Boswell "believing the plant to have nothing to do with L. 
complanatum" (in litt. to Druce, 3 June 1883). It is numbered 1834* and obviously is 
meant to be bound next to 1834 (L. alpinum); no description was given in the text by 
Boswell-Syme. Druce rectified this in a valid publication (1892) distinguishing it from L. 
alpinum "by its larger size, by its flattened spreading branches, with central leaves on the 
flattened stem more erect than the lateral", giving "Scotland, Westmoreland, Cumberland, 
Gloster and Worcester" as localities. Tab. 1834* of Boswell-Syme is referred to and hence 
the type is Reader's plant from Woodchester. Specimens were distributed widely by Father 
Reader; that in BM is designated here as lectotype with isolectotypes seen from BRIST, 
CGE, K and OXF. They all possess cones although mostly over-mature, but are a very 
good match for continental D. issleri. Edward Marshall compared the Woodchester plant 
with material labelled L. complanatum in the Linnean herbarium and published (1891) 
an opinion that Reader's plant "must go under L. alpinum L." and "L. complanatum 
ought to disappear from our list"; this is understandable, the plant is much closer to alpinum. 

As for other material recorded as L. complanatum (or L. alpinum var. decipiens), 
Babington (1883) referred also to Skye and Ross. The Skye plant was collected by Prof. 
Marmaduke Alexander Lawson and H.E. Fox in 1868 and was exhibited at the Linnean 
Society on 22 November 1885 by J.G. Baker. A comment in the Gardeners' Chronicle 
(Anon. 1886) states that "minds have been set at rest by the fine specimen of this species 
[L. complanatum] from the Somerset side of Exmoor", again exhibited by J.G. Baker at 
the Linnean Society on 17 December, 1885. Both are potentially D. issleri (v.i.). Other 
specimens from Scotland, including ones collected by William Gardner on the Sidlaw Hills, 
and by Druce and E.S. Marshall in N. Scotland (e.g. Ben Avon, Banff, v.-c. 94, and Lochnagar, 
c. 3000 ft [1000m,]v.-c. 92) were subsequently labelled var. decipiens, as were many more 
at that period; all have proved to be etiolated forms of D. alpinum. Similarly the specimen 
collected near Advie, Grantown-on-Spey, J.S. Gamble, 1871 (K) is recorded by McCallum 
Webster (1978) as var. decipiens. All those specimens seen are sterile and are luxuriant 
or etiolated forms (through being in dense herbage) of D. alpinum (see below), although 
the Lochnagar material (and site) warrants further investigation. Druce (1916) after 
summarising the history of L. complanatum s.l. and its appearance in Britain, also recorded 
the closely related species Lycopodium chamaecyparissus A.Br, from Ingleborough. The 
specimen on which that record is based, at Manchester Museum (MANCH), is D. alpinum, 
but on the same sheet is a good specimen of L. chamaecyparissus (= D. complanatum 
subsp. chamaecyparissus (A. Br. ex Mutel) Kukk.) obviously misplaced in mounting and 
a suspected source of the error. 

TAXONOMIC CONCEPTS OF L. COMPLANATUM COMPLEX 
It is clear from what has been said above that the status of the names to be applied to 
these records was by no means clear-cut, even if differences were distinct. Those botanists 
who looked beyond our shores realised there were other taxa in Europe and North America 
which might be overlooked. Sir Joseph Dalton Hooker (1884) was one who did not go 
into such problems in depth; he regarded L. alpinum and L. complanatum as subspecies 
- a plausible view - and placed the Gloucester and Worcester plants in ssp. complanatum. 
The status was even reduced further in Bentham & Hooker (1887) where it is dismissed 
as a form *"of warmer climates". As development and change of land-use destroyed lowland 
heaths of Britain, the true identity of these clubmosses became an academic study. 

Early 20th century botanists in Europe reviewed the group from time to time for selected 



DIPHASIASTRUM ISSLERI 



259 




K 



^. ^v ~— * yi^ J~fc„~2 7?'. A *— *. 

-€.*. p.im .V* tCM. .11 f1 3) 









X- 3k4**«' 



7WW 



693 




FIGURE 1. Lectotype specimens of Lycopodium alpinum race issleri Rouy in Issler's Herbarium 
(BASBG). (Photo: Botanisches Institut der Universitat, Basel.) 



260 FERN GAZETTE: VOLUME 13 PART 5 (1989) 



areas but their work did not impinge on the British scene. Then, during the 1950's, a 
Belgian botanist, Dr Andre Lawalree, looked at those forms which were intermediate between 
L. alpinum and L. complanatum, initially in the Low Countries. He described (1957) as 
a species, L. issleri (Rouy) Lawalree based on the L. alpinum variety (actually "Race") 
described by Rouy (1913) from material collected by E. Issler in the Vosges mountains, 
Alsace. Figure 1 shows a sheet from the Issler Herbarium, now at the Botanisches Institut 
Basel (BASBG). Lawalree spread his interest to identifying material in British herbaria, 
and around that time his determinations were sent up by vice-county recorders to BSBI 
Proceedings. Lawalree's determinations included specimens from Llynn Idwal (in BM, NMW 
and OXF), already listed by Druce as L. a. var. decipiens, as L. issleri, and this was taken 
up by Harrison (in Hyde, Wade & Harrison 1969). Lawalree's concept unfortunately included 
etiolated specimens of D. alpinum, and those from Wales, in the opinion of this author, 
were that species. 

Following this account, Joan Wilce, when a student at Michigan University, looked 
at the 'Complanata Section' of Lycopodium (later to be segregated and accepted as a genus, 
Diphasiastrum) for a Ph.D. project. She suggested (1965) that L. issleri was a hybrid, mainly 
on grounds of partial spore abortion, with L. alpinum as one parent and a member of 
the L. complanatum group as the other. This work stimulated more local appraisals in 
Europe, and two are substantial and worthy of mention of mention here. 

First is that of Ilka Kukkonen (1967) who, based in Finland, orientated his work on 
northern and central European material and did not discuss the D. issleri problem to any 
great extent. Nevertheless his discussion of the characters of the D. complanatum complex 
is pertinent as is his account of hybridity within the complex and the ecology of the species 
in Finland. 

The second publication on this group is by Anna Pacyna (1972a, b) who studied Polish 
material, including 15 specimens of D. issleri. Her biometrical and statistical account is 
substantial and she concludes that "D. issleri ought to be considered a species of hybrid 
origin". It is interesting that although D. complanatum is a common plant in the lowlands 
throughout Poland and seldom encountered In the mountains, and then very rarely to 1200m 
altitude, D. issleri is recorded in the same altitudinal zone as D. alpinum (700 to 1200 
or rarely 1600 m). 

I have found signs of spore abortion in both the Gloucester specimens and that recently 
found in the Malvern Hills and have hitherto treated the taxon as a hybrid (e.g. in Jermy 
et al. 1978), whilst accepting that fertile spores are formed and can establish new populations 
where conditions are suitable. However, as D. complanatum sensu stricto has never been 
proven for Britain it seems unlikely that the hybrid has been formed de novo. In view 
of the fact that issleri is intermediate between alpinum and complanatum and that partly 
fertile hybrids appear to occur, the opinion of Domin (1937), that these taxa should be 
treated as subspecies, has a lot of merit. Furthermore, Kukkonen (1967, 1984) presented 
good evidence for this concept in other European taxa of the complex. I therefore propose 
that they are accepted at that rank in the British List (synonymy for Europe in Derrick, 
Jermy & Paul 1987) as follows: 

Diphasiastrum complanatum (L.) J. Holub, Preslia 47: 108 (1975) subsp. complanatum, 

Type: Herb. Linn. 1257. 20 (LINN). 
D. complanatum subsp. issleri (Rouy) Jermy comb. nov. 

Basionym: Lycopodium alpinum L. race issleri Rouy, Fl. France 14:489 (1913). 

Lectotype: France, Vosges Mts, on the central ridge between the pass and the Reisberg, 

on granite, 1298m, E. Issler, 13 Aug 1908 (BASBG). 
D. complanatum subsp. alpinum (L.) Jermy comb.nov. 

Basionym: Lycopodium alpinum L., Sp.PL: 1104 (1753). 

Type: Herb. Linn. 1257.15, Lapponia (LINN). 



DIPHASIASTRUM ISSLERI 



261 




FIGURE 2. Silhouette of Diphasiastrum complanatum subsp. issleri from Canisp, West Sutherland. 
(A.G. Kenneth, 1985, BM). 




A 



B 



FIGURE 3. Diphasiastrum complanatum leaves from second-year-old branches, seen from below. 
A. subsp. complanatum B. subsp. issleri. C. subsp. alpinum. 



262 FERN GAZETTE: VOLUME 13 PART 5 (1989) 



IDENTIFICATION OF DIPHASIASTRUM COMPLANATUM SUBSP. ISSLERI 
The morphology of issleri is intermediate between alpinum and complanatum. If the plants 
are fruiting then the elongated peduncles of the often branched strobilus are distinctive, 
as is the more open, flabellate sterile branch pattern, with the branch angle quite wide. 
The compressed appearance of the lower leafy branch of complanatum is seen in issleri; 
in alpinum the leaves are laxer and the branch appears more terete. Dostal (1984) illustrates 
this well for European material, and Page (1982) gives a figure of the Gloucester plant; 
a silhouette of the Canisp material is given in Fig. 2. In alpinum the alternate branches 
tend to lie in one plane, but in issleri the branches leave the main stem more or less in 
a whorled arrangement, giving a rigid look which, with a somewhat more grass- or yellow- 
green colour (i.e. less glaucous) makes the plant stand out in the community. 

Dostal (1984) also gives the spore size for complanatum (30-38 um); that for alpinum 
is 42-47 um. D. Tennant sent material of the Canisp plant to R.H. Roberts who on measuring 
the spores found them closer to alpinum (mean 41.8 um; Tennant, pers. comm.), comparing 
well with the author's measurements of the Woodchester plant. It is interesting that Roberts 
found the spores of an untypical form of alpinum (sent to him also by Tennant, v.i.) from 
about 300m above Kenneth's site, to be even smaller (mean 35 um). On other morphological 
characters those plants fall within the variation of that species. 

The shape of the sporophyll in the three taxa can be seen to be subtly different. In 
complanatum it is very broadly ovate, almost orbicular with an apiculate tip; in alpinum 
it is more narrowly ovate with an attenuate tip; in issleri it is intermediate. However, there 
is considerable variation throughout the length of the cone, and because most material 
found is sterile, I prefer to emphasise the vegetative characters. 

The shape and size of the leaves on branches of the previous year's growth, are the 
best characters to consider when one is presented with a few branchlets only. Those leaves 
facing to the ground (so-called ventral leaves) on complanatum are narrowly subulate- 
triangular, adpressed to the stem for most of their length; those of alpinum are lanceolate- 
ovate, with an obtuse apex, or more often trowel-shaped (trullate) with a thick petiole 
forming the right-angle of the trowel handle so that the leaf stands away from, and with 
the blade parallel to, the stem, although on drying or pressing, the blade is often 
characteristically skewed; and those of issleri are intermediate in shape, being in British 
material narrowly triangular with an acute apex and lacking the angled petiolate base. 

The shape of the lateral leaves on these older stems is a useful character and has not 
been commented on before, and is in my opinion very helpful in diagnosing issleri. In 
complanatum (Fig. 3A) the keel of the leaf descends parallel to the stem and is abruptly 
rounded as it approaches the leaf below. The leaf of alpinum (Fig. 3C) rapidly narrows 
to the base, forming a slender 'waist', when seen from the flattened sides, just above the 
insertion of the next lower pair of leaves; issleri (Fig. 3B) shows the intermediate state, 
the keel not being parallel, but truncate below, rounding abruptly to the stem. Also in 
complanatum the free portion of the lateral leaf is about one third (or even less) of the 
lower fused base; in alpinum the free portion may be almost half the total length. In British 
issleri the free portion is approximately one third. 

Branches of alpinum when growing in deep litter (or in dense Cladonia lichen turf) 
often become etiolated and appear flattened as in complanatum. Such forms have been 
referred to the latter species in error in British records and herbaria. Close examination 
shows the ventral leaf to he trullate and petiolate, and the distinct taper to the narrow 
waist even more obvious. 

RECORDS OF D. COMPLANATUM SUBSP. ISSLERI 
The most recent records published of this taxon (as D. x issleri) are by Jermy et al. (1978). 
In the light of this recent reassessment the distribution shown there must be revised. Whatever 



DIPHASIASTRUM ISSLERI 263 



its taxonomic status, this taxon is very rare in the British Isles and should not be collected 
except to verify its identity if field observation and photography prove inadequate. The 
distribution of issleri clearly falls into two categories: a lowland south and eastern England 
ecotype of which the Woodchester plant is clearly the best example. Other records are 
based on small samples of plants in herbaria (except for the Broad Down, Malvern, plant) 
but are, in my opinion, this ecotype. The second category is the Scottish low mountain 
populations, directly comparable to the Continental material and associated with areas of 
ancient pine forest. Records of doubtful identity are given in square brackets. 

ENGLAND AND WALES 

[V.-cs 4/5 North Devon/S. Somerset: Specimens from "Exmoor" in BM with no date or 

collector are sterile but possibly this taxon and may be the gathering reported in the Gardeners' 

Chronicle (Anon. 1886) as being similar to Professor Lawson's Skye plant. A scrappy specimen 

from "near Lynton" (W.R. Lawson, Dec. 1885) in BM may be the same gathering and 

may also be issleri. Typical alpinum has also been recorded from the area however. 'No 

post- 1930 records of either taxa have been established]. 

V.-c. 12 North Hants.: Lower Waggoner's Wells, Bramshott, specimen comm. C.C. Babington 

in BM and CGE. The plant has not been recorded there this century but much heath 

remained into the beginning of this century and indeed broom-making from the heather 

continued well into the 1930's. Earlier this century Bramshott Common was grazed by 

cattle and this (or the increased nitrogen) would not have improved the chances of the 

clubmoss. 

V.-c. 34 West Glos.: near Woodchester, Stroud, H.P. Reader, several specimens collected 

over the years 1881-1884 in BM, BRIST, CGE, GL, K, LIV, NMW and OXF. 

"Gloucestershire" but no locality given (but presumably from near Woodchester) coll. ? 

Evans, 10 July 1885 (E). 

Reader, whose activity played a significant part in Gloucestershire botany whilst he 
was incumbent at Woodchester found the specimen in "one of the many valleys which 
intersect the Cotswolds about Stroud, where the ground is broken up into several ferny 
knolls, divided by streamlets". Upon the side of one of those knolls was the Lycopodium. 
The oolite disappears at that point "leaving the lias exposed, or but thinly and partially 
covered by a strip of Fuller's Earth". "In consequence", Reader says in a letter to Druce, 
"the flora of this little tract (some six acres in extent), differs widely from that of the 
surrounding country and includes a large number of plants which ... are not found, or 
found but rarely, on the Cotswolds". He mentions a number of species including Lycopodium 
clavatum, Oreopteris limbosperma, Calamagrostis epigeios, Carex pallescens, C. pilulifera, 
Gnaphalium sylvaticum and Vaccinium myrtillus, the mosses Plagiothecium denticulatum, 
Pleurozia schreberi and Polytrichum juniperinum, and the lichens Baeomyces rufus, Cladonia 
cervicornis and C. digitata. As far as I can discover, Reader at no time recorded the exact 
locality but an annotation in Augustus Ley's copy of Babington's Manual in BM quotes 
Woodchester Park, and the geology suggests the south or east sides of Brown Hill. Extensive 
searching by the author and Sonia Holland CBSBI Recorder for Gloucestershire) has failed 
to produce either the plant or a likely habitat. 

V.-c. 37 Worcester: Hartlebury Common, Kidderminster, Churchill Babington, July 1837 
(discussed above), specimen in CGE, not apparently refound, nor collected since. 

[Great Malvern, Worcestershire Beacon. August 1893, Freeman-Roper (Amphlett & Rea, 
1909 sub L. alpinum). Exact locality not known and no herbarium material has been seen.] 

[Little Malvern, Herefordshire Beacon, W.W. Boucher, 20 Sept 1934. (Rea annot. in 
Amphlett &, Rea). In 1980 John Day (pers. comm.) discovered a Diphasiastrum in same 
area (on Broad Down) at 250m alt., in heathland referable to the association Calluno- 
Deschampsietum where Calluna and Deschampsia flexuosa are co-dominant with Potentilla 



264 FERN GAZETTE: VOLUME 13 PART 5 (1989) 



erecta and Vaccinium myrtillus common. Frequent in the area, but as yet not closely associated 
with the clubmoss, are Sorbus aucuparia and Betula seedlings and Pteridium. The low 
herb turf contains occasional Galium saxatile and Luzula multiflora with the following 
cryptogams: Campylopus fragilis, C. introflexus, Cladonia arbuscula, C. ciliata var. tenuis, 
C. floerkiana, C. chlorophaea, Dicranum scoparium and Pleurozium schreberi. 

The site is being monitored by the Nature Conservancy Council and the Worcestershire 
Naturalists' Trust (N. King pers. comm.) and the encroaching birch controlled, with full 
cooperation of the owners, the Malvern Hills Conservators. In November 1986, one plant 
of Diphasiastrum was removed in a turf, so as to keep the rhizosphere intact, and transplanted 
in a nearby site protected from sheep grazing, but not unfortunately from rabbits, the 
effects of which were considerable. No trace of the plant could be found in the Spring 
of 1988. I initially determined this as D x issleri (i.e. D. complanatum subsp. issleri) but 
in the light of further experience must regard it as a population of subsp. alpinum which 
in the past has hybridized with subsp. issleri. It is morphologically half way between issleri 
and alpinum: it has the leaf structure of the former, and the branching, colour and shorter 
cone peduncle of the latter. Such populations should not be unexpected. The taxon issleri 
is most certainly of hybrid origin (v.s. ) but the balance of genes (and therefore characters) 
will not always be identical, and populations will drift towards one or other parent.] 

SCOTLAND 

V.-c. 88 Mid Perth: Glen Lochy, Allt dubh Ghalair, 975 m. alt., amongst long heather, 

29 July 1890, E.S. Marshall (BM, CGE, SLBI). Same locality and collector, 21 July 1906 

(NMW). 

V.-c. 92 South Aberdeen: Glen Derry, near Braemar, 26 July 1896, F.J. Hanbury (BM); 

there are remnants of ancient pine forests in this valley and the site could have issleri 

still present. 

Strath Nethy, across track to Bynack Beg, in open moorland, between 750 and 800m 
alt., August 1981, D.J. Tennant (E); this is a small sterile branch but a good match for 
the taxon. 

Lochnagar, on the S.E. slopes at 960 m alt., in short heather c. 7-8 ins high, 1961/ 
62, L. Bentley (comm. F. Rose) (BM). 

Glen Quoich, on eastern slopes of A' Chioch, Beinn a 1 Bhuird, at c. 700 m alt; amongst 
and beneath dwarf heath of Calluna-Vaccinium c. 40 cm high, on shallow peaty skeletal 
soils, 6 July 1988, H. Noltie (E); same locality, 18 July 1988, A.C. Jermy No. 18075 (BM, 
NMW). Subsp. alpinum not associated but within 100 m, on more open alpine heath. 
V.-c. 96 Easterness: S.W. slopes of Geal Charn, Glen Feshie, 2700 ft [810m], on open heather 
moorland on granitic soils, J.A. Wheldon and A. Wilson, July 1909 (BM, BRIST, CGE, 
E, OXF, NMW). Searched for many time since unsuccessfully, but could still be there. 
V.-c. 100 Arran: Glen Easan Biorach, at about 1000 ft [300m] altitude, A. Somerville, 2 
July 1895 (E, OXF). Not collected since. 

Lochranza, Terras herb., 1884 (E). Most likely the same locality as above. Not seen 
since, but this or these could well be there still. 

V.-c. 104 North Ebudes: Skye, "Cumins", M.A. Lawson, July 1868 (BM, CGE, OXF). Nothing 
approaching this taxon has since been collected on Skye but obviously there is great potential 
for it to be there. 

V.-c. 108 West Sutherland: -Canisp, on lower slopes at c. 270m alt., amongst boulders in 
dwarf heath vegetation, July 1985, A.G. Kenneth (BM, E); July 1987, D.J. Tennant & 
A.G. Kenneth (BM). On a mountain with much alpinum, the issleri population was distinct 
in the colour and rigidity of the vegetative branches and showed sporing branches with 
long-pedunculate cones, often branching at their bases (see Fig. 2). An extensive survey 
made of the site, and both around and above it, by Archie Kenneth and David Tennant 






DIPHASIASTRUM ISSLERI 265 



(pers. comm.) in 1987, revealed variation in the alpinum populations, which suggests possible 
interbreeding. The area has much potential for further research and may well hold 
complanatum itself. 

ACKNOWLEDGEMENTS 
I am grateful to the late Archie Kenneth whose friendship I had enjoyed for many years 
and who re-kindled my interest in this problem through finding such an excellent site in 
Sutherland, the discovery of which was a further example of his astute field observation. 
I am also indebted to David Tennant who has kindly read this manuscript, and whose 
acute observations on the Canisp plants were freely given to me and have been incorporated 
into this paper. I am also grateful to Dick Roberts for allowing me to quote his spore 
measurements, and to Noel King for his notes on the Worcestershire plant and site. 

I would like to thank the Director of the Botanisches Institut der Universitat Basel, 
and Drs Christian J. Heitz and Tadeus Reichstein, for providing photographs of Issler's 
specimens held in the Basler Botanische Gesellschaft herbarium. The Editors of the Gazette 
made constructive comments on this paper for which I am grateful. 

REFERENCES 
AMPHLETT, J. & REA, C. 1909. The Botany of Worcestershire, Birmingham. 
ANON. 1886. Lycopodium complanatum. Gardeners' Chronicle 1886: 18. 
BABINGTON, C.C. 1867. Manual of British Botany (ed. 6). London. 
BABINGTON, C.C. 1874. Manual of British Botany (ed. 7). London. 
BABINGTON, C.C. 1881. Manual of British Botany (ed. 8). London. 
BABINGTON, C.C. 1883. [Manual of British Botany] Additions and corrections 2. London. 
BENTHAM, J.G. & HOOKER, J.D. 1887. Handbook of the British Flora. London. 
BOSWELL-SYME, J.T. 1886. English Botany (ed. 3) 12: tab 1834*. 
DERRICK, L.N., JERMY, A.C. & PAUL A.M. 1987. Checklist of European pteridophytes. 

Sommerfeltia 6: 1-95. 
DOMIN, K. 1937. On Lycopodium issleriRouy in Czechoslovakia and on the variability of our Lycopodia 

of the Section Heterophylla Spring. Bull. Int. Akad Tcheque Sci. 38: 131-137. 
DOSTAL, J. 1984 Genus Diphasiastrum Holub in Kramer, K.U. (ed.) Pteridophyta in Hegi, G. Illustrierte 

Flora von Mittel-europa, I. Berlin & Hamburg. 
DRUCE, G.C. 1882. On Lycopodium complanatum L. as a British plant. J. Bot. London. 20: 321- 

323, tab 233. 
DRUCE, G.C. 1892. Plants of Glen Spean, Westerness. Ann. Scot. Nat. Hist. 1892: 182-185. 
DRUCE, G.C. 1916. Plant notes, etc. for 1915. Bot. Soc. Exch. Club Br. Isles 4 (3): 219-223. 
GIBBONS, E.J. K1975. The Flora of Lincolnshire. Lincoln. 
HOOKER, J.D. 1884. Students' Flora, ed. 3. London. 
HYDE, H.A., WADE, A.E. & HARRISON, S.G. 1969. Welsh Ferns, Clubmosses, Quillworts and 

Horsetails. Cardiff. 
JERMY, A.C, PERRING, F.H., FARRELL, L. & ARNOLD, H. 1978. Atlas of Ferns. London. 
KENT, D.H. & ALLEN, D.E. 1984. British and Irish Herbaria. London. 
KUKKONEN, I. 1967. Studies on the variability of Diphasium (Lycopodium) complanatum. Ann. 

Bot. Fenn. 4:441-470. 
KUKKONEN, I. 1984. Nomenclature combinations of Finnish vascular plants. Ibid. 21: 209-21 1. 
LAWALREE, A. 1957. Un Lycopode ardennais meconnu, Lycopodium issleri. Bull. Soc. Roy. Bot. 

Belgique90: 109-120. 
LEES, E. 1867. The Botany of Worcestershire. Worcester. 

LLOYD, J. 1867a. Lycopodium alpinum in Hampshire. Gardeners' Chronicle 1867: 808-809. 
LLOYD, J. 1867b. [Correspondence]. Gardeners' Chronicle 1867:997. 
MARSHALL, E.S. 1891. Short Notes: Lycopodium complanatum L. J. Bot. Lond. 29: 186. 
McCALLUM WEBSTER, M. 1978. Flora of Moray, Nairn and East Inverness. Aberdeen. 
PACYNA, A. 1972a. Biometrics and taxonomy of the Polish species of the genus Diphasium Presl. 

Fragm. Flor. Geobot. 18: 255-297. 
PACYNA, A. 1972b. Distribution of the genus Diphasium Presl in Poland. Ibid. 18: 309-341. 
PAGE, C.N., 1982. The ferns of Britain and Ireland. Cambridge. 
ROUY, G.C.C. 1913. Fl. France 14: 489. Paris & Rochefort. 
WATSON, H.C. 1870. A Compendium of the Cybele Britannica. London. 
WILCE, J.H. 1965. Section Complanata of the genus Lycopodium. Beih. Nova Hedw. 19: 1-233. 



266 FERN GAZETTE: VOLUME 13 PART 5 (1989) 



REVIEW 

PTERIDOPHYTE FLORA OF OAXACA, MEXICO by John T. Mickel and 
Joseph M. Beitel. 568 pp. Memoirs of the New York Botanical Garden volume 
46. 1988. ISBN 89327 323 6. Price $94.85 for U.S. Orders, $96.80 for non U.S. 
Orders. 

The Introduction to this book outlines the development of fern floristics for Mexico over 
the past 150 years. From 182 species reported by Martens and Galeotti in 1842, we learn 
that there are now 690 species known in Oaxaca alone out of a present estimate of 1000 
or more for the whole country. 

The complex physiography of this State has much to do with patchy fern collecting 
in the past, as well as the wide natural taxonomic diversity of the fern flora recently revealed. 
Moreover, the range of ecology and the discrepancies in the known flora between this 
and neighbouring States, suggest that primary exploration still has much to yield. 

Oaxaca is phytogeographically at a crossroads between the Caribbean elements of the 
wet windward slopes of the mountain ridges, the arid 'cheilanthoid' flora stretching from 
the north and Central Mexico, and the tropical Central and South American components 
extending to their northernmost limits. 

The largest genera, Asplenium (55 spp.), Thelypteris (45 spp.), Cheilanthes (44 spp.), 
Selaginella (39 spp.), Polypodium (37 spp.) and Elaphoglossum (33 spp.), (nearly 37% of 
the flora in 6 of the 101 genera in 27 families represented) demonstrate clearly, in whole 
or in their parts, the varied ecological characteristics of this fascinating region. 

Descriptions of species are mostly concise and adequate, being sometimes shortened 
and diagnostic for rare or closely related taxa. Explanatory notes are frequent especially 
for new names of which there are about seventy including new species (described here 
for the first time), hybrids, some varieties and, happily, no subspecies. Groups of closely 
related species often have a note following one member of the group entailing cross-reference 
and much turning of pages because of the alphabetical sequence. The book has alphabetical 
arrangement of genera and species throughout. This leads to a comment on the choice, 
quite clearly made by the authors, to 'ignore families' because 'utility should be the prevailing 
guide in selecting an arrangement'. 

But what is utility? An alphabetical catalogue is useful for someone who wants to establish 
quickly a fact or two about a plant of which the name is already known. 

Illustrations, especially first-class fully vouchered line-drawings, like the 129 plates at 
the end of this book, are useful imperatives in providing visual affirmations of appearances 
and relationships. Here, they are arranged according to a cryptic conspectus of 27 families 
at the beginning of the book. What happens in between has nothing to do with this or 
any other system. 

Utility is in no way better served than in systems of classification. They have the purpose 
of summarizing' the characters of groups within groups as a shorthand to learning and 
research in intellectual processes. The foundation of their intrinsic merit, a normal expectation 
in a Flora, is totally lost in the cheerless formality of the alphabet. 

Regrettably, there are precedents for this, and even more regrettably they come from 
the professional eminence of the Flora-writing fraternity. There is, as our authors state, 
'widespread disagreement among pteridologists regarding familial and generic phylogenetic 
arrangements (of ferns).' But has this book been prepared so diligently and painstakingly 
for pteridologists only? How many aspiring pteridologists are there in the State of Oaxaca? 
Are the shorthands and mnemonics of botanical systematics irrelevant to students and other 
interested people, even other botanists, in Mexico and elsewhere? Of the 27 families in 
the 'cryptic conspectus', 17 are quite indisputable in their circumscription. The other 10 

Contd. on p. 276 



FERN GAZ. 13(5) 1989 267 



COMPRESSION AND SLINGSHOT MEGASPORE EJECTION IN 

SELAGINELLA SELAGINOIDES - A NEW PHENOMENON IN 

PTERIDOPHYTES 

C.N. PAGE 
Royal Botanic Garden, Edinburgh EH3 SLR, Scotland 

ABSTRACT 
Observation of the megaspore and microspore dispersal mechanism of Selaginella 
selaginoides shows that the release of the two types of spores takes place by very different 
methods. 

Microspore discharge takes place passively, while megaspore dispersal takes place 
by an active discharge mechanism, termed here 'compression and slingshot ejection'. 

No active megaspore ejection has, as far as I am aware, been hitherto reported in 
this or any other heterosporous pteridophyte genus. 

This mechanism is discussed and interpreted in relation to the outlying taxonomy, 
ecology and palaeobotanic history of Selaginella and is suggested to be a primitive and 
perhaps ancient evolutionary trait within the genus. 

INTRODUCTION 

Selaginella selaginoides (L.) Link is a small, moss-like plant, which occurs widely amongst 
low-growing vegetation throughout a circum-boreal range (Polunin 1959, Page 1988). It 
is also one of the very few pteridophytes in the native flora of Britain and Ireland which 
is heterosporous. Each cone, which matures in Scotland by mid- to late August or early 
September, usually bears an assortment of microsporangia and megasporangia in the axils 
of its weakly-differentiated, leaf-like sporophylls. Sometimes sporangia of each type occur 
somewhat randomly within a cone. More often, microsporangia and megasporangia are 
zoned within each cone, with the megasporangia always at the base, and sometimes there 
are multiple zones of micro- and megasporangia, especially in larger cones. At maturity, 
the simple, globose microsporangia each liberate a large number of small, orange microspores; 
the 4-lobed megasporangia each liberate only four large, white megaspores. Microspore 
discharge takes place passively, while megaspore dispersal takes place by an active discharge 
mechanism, termed here 'compression and slingshot ejection'. 

Although an active microspore-dispersal mechanism has recently been reported in some 
tropical Selaginella species (Koller & Scheckler 1986), no active ejection of megaspores 
has, as far as I am aware, been hitherto reported in this or any other heterosporous 
pteridophyte genus. 

This mechanism is discussed and interpreted in relation to the outlying ecologic, taxonomic 
and palaeobotanic position of S. selaginoides, and is suggested to be a primitive and perhaps 
ancient evolutionary trait within the genus. 

MATERIALS AND METHODS 
The phenomenon of sudden, ballistic, megaspore ejection in S. selaginoides was initially 
discovered by the author by simple hand-lens observation of in situ specimens in the field 
in East Perthshire, Scotland. Subsequent further in situ observations made on more than 
50 wild, mature specimens showed it, indeed, to be a general phenomenon of the species 
throughout its range in Highland Scotland. 

Supplementary laboratory observations were made on isolated cone material transferred 
when freshly gathered in August 1987 and August 1988 to laboratory environments at 
Kindrogan Field Studies Centre, Perthshire and to the Royal Botanic Garden, Edinburgh, 
Scotland, respectively, enabling the exact sequence of events to be more precisely studied. 
Megaspore discharge distances quoted were those achieved in a laboratory atmosphere, 
measuring from the position of the cone to the final settlement point of the megaspore 
from a still atmosphere on a smooth dark surface. 



268 



FERN GAZETTE: VOLUME 13 PART 5 (1989) 





Fig 1, a - b 'going, going, gone!'; a: Megasporangium walls fully withdrawn from around a quartet 
of megaspores, to the point of being fully primed, with the megaspores on the point of sudden and 
violent release; b: the moment immediately following discharge, with the now-empty megaspore cradles 
formed from the original megasporangium wall showing the large upper and lower lobes and the 
much smaller lateral ones (x 64). Photos: C.N. Page. 



OBSERVATIONS 
Compression and slingshot megaspore ejection 

Each microspore of S. selaginoides is c. 30 um diameter, each megaspore c. 690 um diameter, 
the megaspores being thus approximately 23 times the diameter of the microspores, and 
over 12,000 times their volume. Indeed each megaspore is thus sufficiently large to be clearly 
seen with the naked eye. SEM details are well illustrated by Tryon (1949: 424). 

Field and laboratory observations on native Scottish material show that although the 



SELAGINELLA MEGASPORE EJECTION 269 



* 1 

* :-o •- | • -o 

# a ) b 




O 



■-0-— : , 

d 



Fig 2, a - d, diagrammatic representation of the megaspore discharge sequence of Selaginella selaginoides 
described here; a - b, FRONTAL VIEW, a: quartet of megaspores with central pair becoming squeezed 
by compression action between the upper and lower megaspores in the axil of a deflexed megasporophyll 
(thick solid arrows indicate main compression force directions); b: central megaspore pair suddenly 
ejected as a result of this compression action; c - d, SIDE VIEW, c: larger arms of megaspore cradle 
carrying the upper and lower megaspores snap forward as an immediate result of loss of the central 
megaspore pair; d: the upper and lower megaspores are hurled violently from the cone by the slingshot 
action resulting (slender solid arrows indicate direction of initial megaspore and cradle movement, 
thin dotted arrows, the ultimate direction of megaspore ejection). 

microspores are dispersed at maturity in a totally passive fashion, megaspore dispersal takes 
place by an active discharge mechanism, termed here 'compression and slingshot ejection'. 
By this mechanism, the megaspores of each ripe megasporangium are shot ballistically from 
the plant, dispersing them to greater distance than would be likely to be achieved by passive 
dispersal alone, as well as through a wide radial settlement pattern. 

This discharge process is always similar and spectacular. It is best analysed in three 
phases: that of megasporophyll priming, that of megaspore ejection, and that of the dispersal 
distance achieved by the resulting megaspore trajectories. 

Megasporophyll priming 

The megaspore discharge process of S. selaginoides involves the once-only (for each 
megasporangium) sudden and virtually explosive release of what appears to be the slowly 
accumulated tension acquired by successive megasporophylls. 

Each megasporangium, with its adjacent megasporophyll, acts as an independent discharge 
system. Tensioning of the system appears to be initiated by the shrinking and eventual 
splitting of the ripe megasporangium walls. As these shrink, so the whole megasporangium 
becomes drawn progressively deeper into the axil of its subtending megasporophyll blade, 
forcing the four megaspores contained within tightly together. This axial retreat of each 
mature megasporangium causes a gradual backward-flexing of the whole subtending 
megasporphyll blade, which consequently becomes forced into an increasingly divergent 
angle from the axis of the cone. As it becomes deflexed, it appears to become increasingly 
tensioned. Indeed, even casual observation of almost any fully mature cone of this species 



270 FERN GAZETTE: VOLUME 13 PART 5 (1989) 



in the field typically shows one or more deflexed megasporophyll blades standing at an 
abruptly divergent angle from those of the rest of the cone, although no significance has, 
as far as I am aware, ever been previously attributed to this. Only megasporophyll blades 
appear to do this, and each only once, when the megasporangium in their axil is fully 
mature and about to dehisce. It appears to be this gradually accumulated tension which 
ultimately powers the whole ejection event. 

The duration of the period of tensioning varies according to the speed of splitting and 
shrinkage of the mature megasporangium wall, and hence with the humidity of the air. 
In a dry laboratory atmosphere, it usually took place fully over the course of about a 
2-minute period. In the field, periods of from 5-30 minutes or more seem more usual. 

As the tensioning of the megasporophyll blade proceeds, so the shrinking megasporangium 
wall, splitting along approximately X-shaped predetermined lines of weakness, separates 
around the spores into four discrete arms. Each resulting arm clasps, but gradually withdraws 
from, around one megaspore, by slowly sliding back to form a cradle around its megaspore's 
periphery. At this stage, the megaspores can be seen to be arranged as two decussate pairs: 
a medial pair, lying in the plane of the midvein and cone axis, and a lateral pair, lying 
perpendicularly to the first pair and withdrawn partly between them (Fig. la). 

Drawn maximally into the angle between the cone axis and the deflexed megasporophyll 
blade, the quartet of megaspores in the fully-tensioned system appears to be subject to 
especially strong compressive forces between the tightening megaspore cradle arms, which 
themselves are compressed within the axil of the megasporophyll blade. The lateral pair 
of megaspores are especially tightly compressed between the two medial megaspores which 
squeeze their sides, and the tips of their megasporangium walls, which still just retain them. 
The megapsores themselves are hard and rigid objects, and appear not to easily distort 
to this compression. Under a low power microscope, isolated megaspores can be seen to 
have walls which are externally ornamented only by low, rounded, projections. But when 
newly revealed by the withdrawing megasporangium wall, the fresh spores in life have a 
smoothly glossy appearance and apparently moist surface. Indeed, as the megaspores become 
progressively more visible, they look like a clutch of four, shining, wet eggs, held in a 
tight and shrinking nest. 

Once fully primed, with the megasporophyll blade deflexed to an angle of about 80° 
from the cone axis, and each megaspore largely revealed, but still held around its periphery 
by the embrace of one arm of the retreating megasporangium wall, the whole system is 
ready to actively eject the megaspore quartet revealed (Fig la). 

Megaspore ejection 

Sudden total release of the slowly accumulated tension occurs when the four mature 

megaspores are shot forcefully from the plant, in two rapidly-successive (ie split-second) 

volleys. 

Compression ejection of the first megasore volley: Discharge of the first megaspore volley 
is initiated when the steadily retreating margins of the splitting megasporangium wall 
eventually lose their last vestiges of grip on one opposite (always lateral) pair of 
megaspores; for the arms of the megaspore cradle retaining this lateral megaspore 
pair are always much smaller than those retaining the central pair (see below), they 
are thus always the first to finally lose their grip. At this critical point, the now tightly 
compressed, round and apparently moist-surfaced, lateral megaspores are suddenly and 
forcefully ejected as a pair from the megasporangium, in a manner analogous to two 
round bars of wet soap being equally suddenly and forcefully ejected from a pair 
of squeezing hands! 






SELAGINELLA MEGASPORE EJECTION 271 



Slingshot ejection of the second megaspore volley: With the sudden loss of the first 
megaspore pair, which had, during priming, been forcing between them, the remaining 
medial pair of megaspores are hurled from the cone by each of their spoon-shaped 
wall portions snapping sharply forward with a rapid slingshot-action, now fully releasing 
the remaining tension between the cone axis and the formerly deflexed megasporophyll 
blade. Discharge of the second megaspore volley is initiated immediately by the loss 
of the first spore pair, and thus always follows in nearly-instant succession; their larger 
pair of cruciate arms of the original wall (see below), forming spoon-shaped, sling- 
like cradles now snap sharply together, hurling the two remaining megaspores from 
the cone as they close. The sudden ejection of this second megaspore pair is thus 
like projecting two wet plums from two facing tablespoons, each held at its handle- 
base and tensioned by pulling back the spoon-tip! (Fig. 2 c-d). 

Once all the megaspores have been ejected, the relative sizes of the two pairs of cruciate 
arms of the original megasporangium wal^ can now be most clearly seen. The lateral pair 
are always the smaller, and the medial pair always the larger and more spoon-shaped. 
(Fig. lb). The former function to retain their megaspore pair only until overcome by increasing 
compressive forces, the latter act as cradles to then eject their megaspore pair by their 
different slingshot action. 

Dispersal trajectories 

The resulting discharge distances of the trajectories achieved for the size of the plant are 
impressive. In a still, dry, laboratory atmosphere, trapping of ejected spores on sticky plates, 
held inverted above erect cones, indicates that megaspore ejection can take place to a vertical 
height of 1 metre or more, and that such ejected megaspores, when allowed to land from 
their trajectories, arrive with a random distribution radially around the parent cone at distances 
of mostly 5-45 cm, with a few reaching over 65 cm distance. 

Air-resistance against a moving body of such low mass must be a potent factor in 
slowing its velocity rapidly. However, the laboratory distances measured are those achieved 
in still air, and take no account of the field-situation in which the whole cones and ejectred 
megaspores of S. selagionoides, as well as its passively-released small microspores, in bleak 
upland wild habitats, are more or less constantly subjected to considerable crosswind forces. 
It may, indeed, be such passing breezes which, under field conditions, are effective in promoting 
the initial desiccation process leading to the megaspore discharge event. Further, such breezes 
not only serve to disperse the passively-released microspores, but also quite actively agitate 
the direction of orientation of the cone itself, thereby probably greatly increasing the total 
radial component of the trajectory of megaspores which become ejected at varying radial 
angles into the passing (and probably turbulent) moving air. 

To imitate wind-oscillation, when cones of S. selaginoides were experiementally held 
in the laboratory at varying angles to the vertical, horizontal megaspore discharge distances 
were found to be greatly increased, and horizontal settlement distances of from 1.5 to over 
2 metres from the cone were regularly achieved, with a few megaspores occasionally being 
shot to horizontal distances of over 3m from the parent cone! 

Cone-agitation by crosswind forces thus probably plays a significant role in increasing 
both the absolute distance and randomness of radial direction in which the megaspores 
are finally hurled. 

DISCUSSION 
Although this 'compression and slingshot' method of spore ejection is a structurally simple 
one, it is impresively effective in hurling the large megaspores of this small-statured species 
over appreciable airborne distances, and through a wide radial range of scatter pattern. 
The actual speed of ejection of the spores from the plant (the equivalent of the "muzzle 



272 FERN GAZETTE: VOLUME 13 PART 5 (1989) 



velocity" of a gun) must be considerable, considering the small megaspore mass and the 
height and distance to which each megaspore is shot. Indeed, impingement of the suddenly- 
ejected spores as two successive volleys can certainly be felt on the face of the close observer. 
It also seems to have aspects of considerable biological and palaeobotanic significance. 

Ballistic aspects 

Various mathematical formulae have been proposed to describe the distribution patterns 
of both passively dispersed spores and the ballistics of spores which are violently ejected, 
the latter especially resulting from observation of fungal mechanisms (see, for example, 
Buller 1909, Gregory 1945, 1951, 1961, Ingold 1965, 1971, Gregory & Stedman 1953, Schrodter 
1960 and Sreeramulu & Ramalingen 1961). 

From the aggregate of such studies, two general principles applicable to the contrasting 
spore types of Selaginella seem to emerge: 

One is that for passively dispersed airborne spores, relatively random downwind dispersal 
patterns may be anticipated, analogous to a smoke plume, beneath which is likely to be 
an approximatley logarithmic decrease in settlement density linearly away from the spore 
source in a unidirectional, narrowly triangular downwind pattern. Absolute dispersal distance 
for any spore is likely to be strongly influenced by a multiplicity of environmental variables. 
For achievement of highest dispersal distance, smallness of size promoting slow settlement 
rates and hence long airborne times, is likely to be .of high selective advantage, and, indeed, 
the microspores of Selaginella would appear to fall well within an effective operational 
size for such anticipated missions. 

By contrast, the second principle to emerge seems to be that the subsequent trajectories 
and settlement patterns of violently-ejected spores are somewhat more mathematically 
predictable (if more complex to calculate), with a generally circular pattern of expected 
settlement patterns (modified to perhaps a more ovate shape by crosswinds) reaching a 
maximum density away from the source at a predictable radial distance. Also by contrast 
with microspore size, for highest dispersal distance and for minimum influence to trajectories 
of the vicissitudes of wind currents and air turbulence, rapid speed of ejection as well as 
relatively large mass would be at a strong selective advantage. The size of the megaspores 
of S. selaginoides certainly appear to be within an adequately massive operational size for 
their very different dispersal mechanism to operate effectively in the field. 

Ecological and biological aspects 

In Scotland, the erect cone-shoots of S. selaginoides normally arise to about 1-4 cm above 
the level of the ground. Its montane habitats, as well as its coastal dune ones, are sites 
which are not, and probably have never been, covered by extensive tree growth in post- 
glacial time. Further, it is very probable that S. selaginoides was once a widespread species 
of early post-glacial habitats, characterised by their open vegetation, before the extensive 
development of tree vegetation in Britain and Ireland (Page 1982, 1988), and, indeed, exposure 
of habitats, appears typical of the plant throughout much of its modern, boreal range. 
Were its megaspores merely passively released, at heights of under 4 cm above ground 
level (and often among other low-growing herbage), their opportunity for more effective 
dispersal by wind currents might well be appreciably limited. Ejected into the air, however, 
not only is more distant dispersal likely to be regularly achieved, but so also is a far better 
radial directional scatter of megaspores through wind-agitation of the cone. 

Biologically, the significance of this process is presumably not only in removing the 
sites of new megaprothallial establishment (and hence the site of origin of new sporophytic 
progeny) well away from the parent plant, but also in increasing the opportunity for 
randomness of fertilisation of the resulting megaprothalli by the antherozoids of independent 
microprothalli deriving from passively-dispersed microspores originating from independent, 
differently-placed, upwind, sporophytes. For only when so-removed, in large part, from 



SELAGINELLA MEGASPORE EJECTION 273 



settlement within the same downwind dispersal zone as that of a sporophyte's own 
microspores, do the biological benefits of heterospory in an airborne environment seem 
likely to become most fully realised. 

Elsewhere in the genus, in a wide range of tropical species of Selaginella examined 
by Roller & Scheckler (1986), no active megaspore ejection mechanism was detected, nor, 
so far as I am aware, has nay been by any other author on Selaginella, and it seems 
possible that this may indicate a real absence of this phenomenon amongst the tropical 
species. In these species, a more active microspore dispersal system, however, exists, and 
as such it would seem that in these taxonomically very different groups of Selaginella (the 
species-poor homophyllous one and the species-rich heterophyllous one), a similar biological 
result has probably been achieved by specialisation within each group in violent dispersal 
of one (opposite) spore type only. For it perhaps matters less, biologically speaking, how 
the two types of spores are actually discharged, so long as they are removed from the 
site of the parent plant and that they are discharged in ways that achieve significantly 
different settlement patterns. 

Taxonomic and palaeobotanic aspects 

Within the genus, S. selaginoides is taxonomically a highly distinctive member, with cylindrical 
cones, homorphic leaves and spiral phyllotaxy, there being only one other living species 
(S. deflexa Brack, of the Hawaiian Islands) with a similar general morphology. The sporangial 
arrangement, with a basal megasporangiate zone and a superior microsporangiate one, is 
also restricted within the genus to this pair of species (Horner & Arnott 1963). These aspects 
and the relatively undifferential foliar trait of S. selaginoides (which also includes very leaf- 
like sporophylls) has been taken to indiciate primitiveness within the genus as a whole 
(eg Bower 1908, 1935: 288, Smith 1938, Thomas 1985, Jermy in press). It thus seems possible 
that this 'compression and slingshot' megaspore ejection mechanism may itself be ancient 
within the genus and as such could have existed in similar form in early members of Selaginella. 

At the family level, Selaginellaceae (the only family of the Selaginellales - Thomas & 
Brack-Hanes 1984) appears to have been separate from Lycopodiaceae since at least the 
Lower Carboniferous. For, by this time, heterospory amongst ligulate clubmosses with radial 
shoot symmetry had already evolved (Bower 1908, 1935, Hirmer 1927, Smith 1938, Boureau 
1967, Stewart 1983, Thomas & Spicer 1987), while there is also evidence of perhaps even 
more primitive ligulate lycopods from as early as the middle Devonian (Grierson & Bonamo 
1979). 

At the genus level and below, heterophyllous and homophyllous groups of Selaginella 
appear to have been separate since at least the Upper Carboniferous (Thomas 1985 & 
pers. comm. 1989), from when, as with the other probably equally ancient pteridophyte 
genus Equisetum (Equisetaceae) (Page 1972), small herbaceous forms of Selaginella may 
well have survived relativley unchanged to the present day (eg Taylor 1981, Thomas 1985). 
Evidence for the early existence of small, S. selaginoides-like forms comes from several 
independent sources. Bower (1908), for example, first drew attention to the work of Benson 
(1907), who compared a small Carboniferous plant of lycopod-like form, Miadesmia 
membranacea, with the habit of living S. selaginoides, although the megasporangial characters 
appear to have been even more advanced (see below). Another Carboniferous fossil, Selaginella 
fraipontiiv/as also clearly a plant of herbaceous form and probably sprawling habit (Schlanker 
& Leisman 1969), while comparisons have also been made of the Carboniferous Selaginellites 
crassicinctus (Leisman 1961) and of Paurodendron (Phillips & Leismar 1966) with the living 
5. selaginoides, underlining further the morphologically primitive status and probable very 
great antiquity of the herbaceous habit and general homomorphic form of living S. 
selaginoides. 

Through the fossil record, with the evolution of heterospory, the number of megaspores 



274 FERN GAZETTE: VOLUME 13 PART 5 (1989) 



per megasporangium has been different in different taxa. Even today, some Selaginella 
still show occasional variation in megasporangial number (e^ Duerden 1929), while the 
fossil record shows many possible past avenues of evolutionary opportunity in terms of 
megaspore-reduction numbers. Some of the Carboniferous taxa of Lycopodites (subsequently 
put into Selaginellites) reported by Bower (1908: 301) had 16 or 24 megaspores per 
megasporangium. However, by as early as the Carboniferous, the present-day number of 
S. selaginoides had already been achieved in Selaginella fraipontii (Schlanker & Leisman 
1969), while this number was modified still further in the arborescent Lycopod Lepidophloios, 
by reduction of the developing tetrads to one and the subsequent abortion of three of 
its members, although in this case, the resulting single mature megaspore is believed to 
have been discharged complete with its sporophyll as an airborne unit (Thomas 1978, Thomas 
& Spicer 1987). The megaspore number was also reduced to one in the herbaceous Miadesmia 
membranacea (Benson 1907, Bower 1908) - see below. 

The data from living S. selaginoides suggests that, for the microspores of this species, 
and presumably for those of the fossil forms, evolutionary selection pressure has probably 
always favoured smallness in size and low mass to maximise airborne dispersal opportunity. 
But for airborne megaspores, once larger than a critical size above which passive wind 
dispersal is less effective, selection pressure would seem likely to have favoured the evolution 
of more massive projectiles, and that of an efficient ejection mechanism, helping ensure 
different spore settlement patterns and thereby maximising opportunity for outbreeding 
between prothalli from different original sporophytes. 

Evidence from the fossil record certainly provides abundant indications of widespread 
trends to reduce the megaspore number per megasporangium, wiiile increasing its size and 
mass. It would be of interest to know whether any cone specimen in the long history of 
these fossil heterosporous clubmosses also showed evidence in the form of occasionally 
deflexed megasporophyll blades, or of discharged megasporangial walls of regular structure, 
indicative of the past presence of active megaspore ejection systems. 

Finally, the strategic advantage of mass in a ballistically-ejected megaspore projectile 
would also open opportunity, should it be of subsequent selective advantage, for increase 
in biological reserves. In this connection, it seems of interest to note that Bower (1908: 
300), although he did not know of the megaspore ejection mechanism described here, 
compared the Carboniferous Miadesmia membranacea with living S. selaginoides, but pointed 
out the principal difference that Miadesmia bore in its megasporangia not four megaspores, 
but what is described by Bower as 'a single, seed-like structure' (on the basis of Benson's 
observation that it germinated in situ, presumably into a megaprothallus) (Benson 1907). 
Once an efficient megaspore ejection mechanism had evolved, it would not be too big 
a step to conjecture such a mechanism then advantaging the further evolutionary step that 
Miadesmia suggests, of reduction to a single megaspore, retained through germination to 
a simple prothallus within the protection of its megasporangium, pollinated in situ by the 
germination of airborne microspores, and the whole fertilised-product then efficiently ejected 
as a simple seed. 

ACKNOWLEDGEMENTS 
I am grateful for discussion of these observations with Dr M. Cousens (University of West 
Florida, Pensacola) and for comments on the manuscript from Professor J. McNeil (Royal 
Botanic Garden, Edinburgh), A.C. Jermy (British Museum, Natural History), Dr B.A. 
Thomas (National Museum of Wales, Cardiff) and Dr T.G. Walker (University of Newcastle 
upon Tyne). 



SELAGINELLA MEGASPORE EJECTION 275 



REFERENCES 

BENSON, M. 1907. Miadesmia membranacea Bertrand; a new palaeozoic lycopod with seed-like 

structure. Proc. Roy. Soc. Ser. B 79: 412>2l. 
BOUREAU, E. (ed.) 1967. Traite de Paleobotanique - Tome 2. Massie et Cie, Paris. 
BOWER, F.O. 1908. The Origin of a Land Flora. MacMillan, London. 
BOWER, F.O. 1935. Primitive Land Plants. London: MacMillan. 
BULLER, A.H.R. 1909. Researches in Fungi. Vol. 1. London. 

DUERDEN, H. 1929. Variations in megaspore number in Selaginella. Ann. Bot. 45:451-459. 
GREGORY, P.H. 1945. The disperson of air-borne spores. Trans. Brit. Mycol. Soc. 45: 249-254. 
GREGORY, P.H. 1951. Deposition of airborne Lycopodium spores on cylinders. Ann. Appl. Biol. 

38: 357-376. 
GREGORY, P.H. 1961. The Microbiology of the Atmosphere. London. 
GREGORY, P.H. & STEDMAN, O.J. 1953. Deposition of air-borne Lycopodium spores on plane 

surfaces. Ann. Appl. Biol. 40: 651-674. 
GRIERSON, J.D. & BONAMO, P.M. 1979. Leclerckia complexa: earliest ligulate lycopod (Middle 

Devonian). Am. J. Bot. 66: 474-476. 
HIRMER, H. 1927. Handbuch der Palaobotanik. Oldenbourg: Berlin. 
HORNER, H.T. & ARNOTT, H.J. 1963. Sporangial arrangement in North American species of 

Selaginella. Bot. Gaz. 724:371-383. 
INGOLD, C.T. 1965. Spore Liberation. Clarendon Press, Oxford. 

INGOLD, C.T. 1971. Fungal Spores. Their Liberation and Dispersal. Clarendon Press, Oxford. 
JERMY, A.C. (in press). Selaginellaceae in Kubitsky, K. et al. (eds), The Families and Genera of 

Vascular Plants. 
KOLLER, A.L. & SCHECKLER, S.E. 1986. Variations in microsporangia and microspore dispersal 

in Selaginella. Am. J. Bot. 73: 1274-1288. 
LEISMAN, G.A. 1961. Further observation on the structure of Selaginellites crassicinctus. Am. J. 

Bot. 48: 224-229. 

PAGE, C.N. 1972. An interpretation of the morphology and evolution of the cone and shoot of Equisetum. 

J. Linn. Soc. Bot. 65: 359-397. 
PAGE, C.N. 1982. The Ferns of Britain and Ireland. Cambridge University Press. 
PAGE, C.N. 1988. Ferns. Their Habitats in the Landscape of Britain and Ireland. Collins New Naturalist 

series. London. 
PHILLIPS, T.L. & LEISMAN, G.A. 1966. Paurodendron, a rhizomorphic lycopod. Am. J. Bot. 53: 

1086-1100. 
POLUNIN, O. 1959. Circumpolar Arctic Flora. Oxford: Clarendon Press. 
SCHLANKER, CM. & LEISMAN, G.A. 1969. The herbaceous Carboniferous lycopod Selaginella 

fraitpontii comb. nov. Bot. Gaz. 130: 35-41. 
SCHRODTER, H. 1960. Dispersal by air and water - the flight and landing, pp. 167-227 in Horsefall, 

J.G. & Dimond, A.E., Plant Pathology: an Advanced Treatise. Vol. 3. New York. 
SMITH, G.M. 1938. Cryptogamic Botany. Vol. II. Bryophytes and Pteridophytes. McGraw-Hill, New 

York. 
SREERAMULU, T. & RAMALINGEN, A. 1961. Experiments in the dispersion of Lycopodium and 

Podaxis spores in the air. Ann. Appl. Biol. 49: 659-670. 
STEWART, W.N. 1983. Paleobotany and the Evolution of Plants. University Press, Cambridge. 
TAYLOR, T.N. 1981. Paleobotany. An Introduction to Fossil Plant Biology. McGraw-Hill, New York. 
THOMAS, B.A. 1978. Carboniferous Lepidodendraceae and Lepidocarpaceae. Bot. Rev. 44: 321-364. 
THOMAS, B.A. 1981. Structural adaptations shown by the Lepidocarpaceae. Rev. Palaeobot. 32: 377- 

388. 
THOMAS, B.A. 1985. Selaginella: the persistent pteridophyte. Pteridologist 1: 59. 
THOMAS, B.A. & BRACK-HANES, S.D. 1984. A new approach to family groupings in the lycophytes. 

Taxon 33: 247-255. 
THOMAS, B.A. & SPICER, R.A. 1987. The Evolution and Paleobiology of Land Plants. Croom 

Helm, London. 
TRYON, A.F. 1949. Spores of the genus Selaginella in North America north of Mexico. Ann. Missouri 

Bot. Gard. 35:413-431. 



276 FERN GAZETTE: VOLUME 13 PART 5 (1989) 



Contd. from p. 266 

are a backlash against the artificially unitary, and actually polphyletic 'Polypodiaceae' of 
more contented days. Here lies the main cause of the disagreement, but the 'Polypodiaceae' 
did no injustice to the uninitiated, and so it must be the professionals who are floundering 
in their own confusion. This should not be a position to retreat from but rather another 
opportunity to test an existing system, or to try yet another new one. There is much more 
information available to-day than there ever was in the past, and there are much easier 
ways of sorting it. 

The indefinite prospect of putting up with Catalogues rather than Floras, while we 
wait for a perfect system agreeable to all, is an undesirable alternative. 

The contents of the book are most comprehensive and are the result of years of meticulous 
research. It is a beautifully produced volume of which the authors and publishers should 
be deservedly proud. 

C. D. ADAMS 



SHORT NOTE 

DRYOPTERIS x FRASER-JENKINSII - A CORRECTION 

The name Dryopteris x fraser-jenkinsii was invalidly published in a recent article (Gibby 
& Widen 1983) because a holotype was not properly indicated, and this is now corrected: 

Dryopteris x fraser-jenkinsii Gibby & Widen, hybrida nova 

Holotypus: ex hort. Chelsea Physic Garden, CPG 2189, 25 August 1981 (BM), originally from 
Spain, Oviedo, beside road from Avilcs to Ribadeo, by bend in road, 1km S. of Canero, E. 
of Luarca, 50m alt., mixed Pinus, Quercus and Castanea forest, with D. affinis (Lowe) Fraser- 
Jenkins, D. dilatata (Hoffm.) A. Gray and D. guanchica Gibby & Jermy, coll. C.R. Fraser- 
Jenkins 4899, 5 June 1976. 
Isotypus: (M). 

Hybrida in morphologia D. a/77n; similis. Pinnulae autem plus dissectae ad costam fissae, segmentis 
rotundatis dentatisque. Sporae pleraeque abortivae, vix 5% ut videtur bene evolutae et haec 
grandes, rugosae eis D. affinis similes sed nonnumquam subspineae vel spinulosae. 

REFERENCES 
GIBBY, M. & WIDEN, C.-J. 1983. Three new Dryopteris hybrids from Spain and the Canary Islands. 
Fern Gaz. 12: 267-270. 

MARY GIBBY 

Department of Botany, British Museum (Natural History), 

Cromwell Road, London SW7 5BD, U.K. 

& C.-J. WIDEN, 

Pharmacy of Maunula, Metsapurontie 23, 

SF 00630 Helsinki 63, Finland. 



FERN GAZ. 13(5) 1989 277 



AN ABERRANT FORM OF EQUISETUM TELMATEIA 
(PTERIDOPHYTA) FROM THE WEST OF IRELAND 

MARGARET R.I. WESTWOOD 

School of Molecular and Biological Sciences, The University, Stirling FK9 4LA, 

Scotland 

ABSTRACT 

A population of Equisetum telmateia in Co. Clare, Ireland, was discovered to have a 
number of shoots which produced a proliferation of small cones. These specimens were 
examined and found to have apparently viable spores. Speculation is made that this 
type of cone production may provide a link to extinct horsetails. 

INTRODUCTION 
During a visit to The Burren, Co Clare, a population of Equisetum telmateia Ehrh was 
found at Clifden Hill (National Grid of Ireland 127189) on 6 June 1988. Several unusual 
fertile shoots were present which bore whorls of small cones. The phenomenon of proliferating 
cones is rare, but has been recorded by Pope (1983) in a population of E. telmateia from 
the Isle of Wight, and in various forms and species by Page (1972). 

SITE 
The aberrant specimens were part of a vigorous population of Equisetum telmateia growing 
along ditches on both sides of a minor road. The E. telmateia was the dominant species 
between the ditches and the hedges behind for a distance of about 1 km. The road was 
about 100 m above Ordnance Datum and was situated on a slope between the acidic recently 
glaciated shale lands and the Carboniferous Limestone of the Burren (Willmot 1979). 
Associated species included the calcicol indicators Polystichum setiferum and Asplenium 
(Phyllitis) scolopendrium. Equisetum palustre was also present. 

SPECIMEN DESCRIPTION 
Morphology 

The aberrant fertile shoots possessed nodal branches and had a mean height of 90cm, 
which was generally 10-20cm smaller than wholly vegetative shoots. Fertile stems had senescing 
apical cones which were similar in appearance to normal E. telmateia cones, and the upper 
brlanches carried small cones in whorls around the nodes (Fig. 1). The proliferating cones 
varied from being well filled and terminating short branches (Fig. 2a), to shrunken vestigial 
cones which had reverted to branches (Fig. 2b), or intermediates between these conditions. 
Some lower branches carried two successive cones, (Fig. 2c). Frequently the vegetative 
branches bore further lateral branches (Fig. 1). It should be noted that some branch-bearing 
stems were found which bore single, fertile apical cones, but lacked any profileration of 
cones around the nodes. 

Spores 

Under a light microscope, sporangia of the smallest cones generally appeared fused and 
shrunken, particularly in cones which had reverted to branches. Sporangia from apical 
and larger, non-apical cones produced some abortive material, but many good spores with 
elaters, chlorophyll and a spherical shape were also present (Fig. 3). The ratio of apparently 
viable spores to abortive material tended to increase with cone size, but variation was also 
found between sporangia within a single cone. 

DISCUSSION 
The Burren population differs from that found in the Isle of Wight in that Pope (1983) 
reported small cones only on the tips of proliferating side shoots, while Burren specimens 



278 



FERN GAZETTE: VOLUME 13 PART 5 (1989) 







5c« 



FIGURE 1. Overall structure of aberrant fertile shoots of Eqmsetum telmateia, Clifden Hill, Co. Clare. 
Note the many small intercalary cones. 



AN ABERRANT EQUISETUM TELMATEIA 



279 




a 




FIGURE 2. Details of cone proliferation in Equisetum telmateia. 

(a) cones from upper nodes with well filled sporangia (x 32). 

(b) vestigial cone reverting to branch (x 160). 

(c) alternation between vegetative and fertile growth on the stem node branches (x 32). 



280 



FERN GAZETTE: VOLUME 13 PART 5 (1989) 




C FIGURE 2(c). 



.■■■'■'"■■ 




WHimi' 



l 




FIGURE 3. Spores from a larger proliferating cone of Equisctum telmatciu, showing apparently good 
morphology (x 2500). 



AN ABERRANT EQUISETUM TELMATEL 28 1 



alternate from cone, to branch, to cone. Mr B. Robson (personal communication), has 
reported that the production of multiple cones occurs annually on the site, therefore the 
specimens are not the result of a single isolated abnormality. 

Page (1972) and Page & Baker (1985) note that Equisetum species have a high degree 
of morphological plasticity, which this find reflects. The proliferation of cones reverting 
to shoots is reminiscent of some horsetails known from the fossil record, eg the Permian 
Phyllotheca deliquescens (Goeppert) Schmalhansen (Boureau 1964: 398), the Upper Permian 
Equisetinostachys grandis Rasskazovo (Boureau 1964: 414) and the Upper Triassic Equisetites 
(Equisetostachys) bracteosus Kon'no (Kon'no 1962; Boureau 1964: 435). Populations of 
this type may indicate that the divergence of extinct and modern horsetails may be less 
than often believed. 

ACKNOWLEDGEMENTS 
I am grateful to Dr C.N. Page and Dr J. Proctor for their help. 

REFERENCES 
BOUREAU, E. (Ed), 1964. Traite de Paleobotanique. 3, Sphenophyta & Noeggerathiophyta. Paris: 

Masson. 
KON'NO, E., 1962. Some species of Neocalamites and Equisetites in Japan and Korea. Sci. Rep. 

Tukoko U. Sendai, Japan (2nd ser. (Geol.)), 5, special volume: 21-49. 
PAGE, C.N., 1972. An interpretation of the morphology and evolution of the cone and shoot of 

Equisetum. Bot. J. Linn. Soc. 65: 35-397. 
PAGE, C.N. & BAKER, M.A., 1985. Ecology and geography of hybridisation in British and Irish 

horsetails. Proc. Roy. Soc. Edinb. 86B: 265-272. 
POPE, C.R., 1983. An aberrant form of Equisetum telmateia from the Isle of Wight. Fern Gaz. 12(5): 

303-4. 
WILLMOT, A., 1979. An ecological survey of the ferns of the Burren, Co. Clare, Eire. Fern Gaz. 

12(1): 9-28. 

REVIEW 

AZOLLA UTILIZATION (Proceedings of the Workshop on Azolla use, Fuzhou, 
Fujian, China, 31 March-5 April 1985). Edited by W.H. Smith and Emerita 
Cervantes. International Rice Research Institute, Manila 1987. 

The cultivation of Azolla with rice is traditional in China. It provides supplementary nitrogen 
to the rice crop, suppresses weeds by shading, and provides food for fish and ducks. It 
can be harvested and fed to pigs and domestic fowls. This is the report of an international 
symposium on the cultivation and use of Azolla. In India this practice is increasing; dried 
cow dung and powdered neem cake are used to encourage growth of the fern, which has 
a high phosphorus requirement. Programmes to encourage the use of Azolla are in being 
in the Philippines, elsewhere in Asia, and in Africa and in South America. There are problems 
to be overcome; the plant does not thrive at high temperatures, and it is subject to a 
number of serious insect pests. In Japan Eichornia is preferred as a rice nitrogen source 
and as animal feed. This report is a useful summary of the present state of knowledge 
in this subject. 

F.H. BRIGHTMAN 



282 FERN GAZ. 13(5) 1989 



A NEW SPECIES OF SELAGINELLA (PTERIDOPHYTA) FROM 
CAMEROON, WEST AFRICA 

NAT. QUANSAH 
WWF Aires Protegees, B.P. 738, Antananarivo 101, Madagascar 

ABSTRACT 

Selaginella serrato-squarrosa Quansah, a new species from Cameroon, is described and 
illustrated. 

INTRODUCTION 

In his account of the genus Selaginella Beauv. in West Africa, Alston (1959) listed 20 species. 
A recent taxonomic revision of the genus in West Africa and Madagascar by the author, 
Quansah (1986), has revealed one new species from Cameroon.: 

Selaginella serrato-squarrosa Quansah, sp. nov. Figure 1 & 2 

Species Selaginella heterophylla Selaginellae squarrosae Baker similis sed ab illea foliis marginibus 
serratis, foliis mediis aristatis longe (arista laminae duplo longiore, strobilis aut omnino 
microsporangiatis aut sporophyllis dorsalibus megasporangiatis microsporangiatisque et 
sporophyllis ventralibus omnino microsporangiatis. 

Typus: CAMEROON, Edea, c. 1500m on moist forest floor, 28 April 1948, Nickles 102 (P, 
holotypus). 

Plants erect/sub-erect; branch-system 3-5 pseudopinnate; rhizophores arising at the axils of primary 
branches and restricted to the basal two-thirds of the plant. Leaves anisophyllous, single-veined; 
ligules up to 0.45 mm long, elongate pedate, occasionally bifid. Lateral leaves asymmetrical, 
sub-subulate to ovate-oblong, up to 5.3 x 1.6mm base sub-attenuate to oblique, apex acute, 
margins of basal third serrate-entire, apical two-thirds entire/sub-entire; amphistomatous; aligular 
surface epidermis with sclerotic cells forming patches and/or bands on lamina. Median leaves 
asymmetrical, lanceolate, up to 4.5 x 1.1mm, base oblique (obtuse-cuneate), apex long aristate 
(aristae up to two times the length of lamina), margins serrate; hypostomatous. Axillary leaves 
symmetrical, ovate to narrowly deltate, up to 5.0 x 1.8mm, base truncate to weakly subcordate, 
apex acute, margins of basal half serrate, apical half entire; hypostomatous; aligular surfasce 
epidermis with sclerotic cells forming patches and/or bands on lamina. Strobili bilateral, 
resupinate, up to 5 mm long, with two sporangial arrangements: (i) cone wholly microsporangiate; 
(ii) with the dorsal side containing both megasporangia and microsporangia randomly arranged 
and the ventral side wholly microsporangiate. Sporophylls dimorphous. Ventral sporophylls 
subpanduriform-ovate, up to 2.4 x 1.0mm, base obtuse, apex acute, margins serrate-entire to 
aculeate-entire, with a complete sporophyll-pteryx at the adaxial surface; amphistomatous; 
sporophyll-pteryx with serrate margin. Dorsal sporophylls lanceolate, up to 1.2 x 0.6 mm, base 
obtuse, apex cuspidate to aristate (aristae up to same length as lamina), margins serrate; 
hypostomatous. Megasporangia ovoid-triangular, with similar-sized spores; megaspores 210- 
(M200)-255um in equatorial diameter, trilete, sub-globose, both proximal and distal surfaces 
rugulose. Microsporangia ellipsoid to roundish; microspores 15-(M19)-28pm in widest area, trilete, 
sub-triangular, both proximal and distal surfaces granulose to foveolate. 

This species is closely allied to S. squarrosa Bak. from which it is distinguished by 
its serrate-margined leaves, very long aristate median leaves (aristae up to twice the length 
of the lamina) and two sporangial arrangements of the strobili. S. squarrosa has leaves 
with entire margins, aristae of median leaves up to 1.5 times the length of the lamina 
and strobili with one sporangial arrangement. 

Other specimen examined: 

GABON. Kkan, NE Mela, Mont de Cristal, Halle & Villiers 4788 (K) 

ACKNOWLEDGEMENTS 
The author is grateful to Dr B.A. Thomas, Botany Department, National Museum of Wales, 
Cardiff, for constant encouragement and Mr A.C. Jermy & Dr. N.K.B. Robson, Botany 



SELAGINELLA SERRATO-SQUARROSA 



283 



Vi'. 




m, 



HERB- INST FR. AFRIQUE NOIRE 






HI-RB. MI'S. PARIS. 
| * HOLOTYPE 

Sdag'nella serrato- squarrosa QuanSah 




CMS 



FIGURE 1. Selaginella serrato-squarrosa Quansah. Holotype (Nickles 102, P) showing general habit. 
Photo: by permission by the Trustees of the British Museum (Natural History). 



284 



FERN GAZETTE: VOLUME 13 PART 5 (1989) 




FIGURE 2. Selaginella serrato-squarrosa. a) lateral leaf; b) median leaf; c) anxillary leaf; d) ventral 
sporophyll; e) dorsal sporophyll; f) ligule. a-e = 1mm scale; f = 0.1mm scale, p = sporophyll-pteryx; 
sp = sclerotic patches; v = veins. 



Department, British Museum (Nat. Hist.) for help with the Latin diagnosis. The main research, 
of which this publication is a part, was supported with a grant from the Ghana Government. 



REFERENCES 
ALSTON, A.H.G. 1959. The ferns and fern-allies of West Tropical Africa. Suppl. to the 

2nd ed. of the Flora of West Tropical Africa. VIII + 89pp. Crown Agents for Overseas 

Government and Administrations, London. 
QUANSAH, N. 1986. A taxonomic revision of the species of Selaginella Beauv. subgenus 

Stachygynandrum (Beauv.) Bak. from West Africa and Madagascar. Ph.D. Thesis, 

University of London (Unpubl.). 



FERN GAZ. 13(5) 1989 285 



NEW FERNS OF MADEIRA 

MARY GIBBY 

Department of Botany, British Museum (Natural History), London SW7 5BD 

& 

J.D. LOVIS 

Department of Plant and Microbial Sciences, University of Canterbury, 

Christchurch, New Zealand 

ABSTRACT 

Three new endemic taxa from Madeira are described, Hymenophyllum maderense Gibby 
& Lovis, Asplenium trichomanes subsp. maderense Gibby & Lovis and Ceterach 
lolegnamense Gibby & Lovis; Polystichum x maderense Johnson and other endemic 
hybrids are discussed. 

In the course of a study of the cytology of the fern flora of Madeira (Manton et al. 1986), 
some new ferns were discovered, and although the cytology of these new taxa has been 
described, they still require formal taxonomic description. In addition, another new hybrid 
has been found during the preparation of the forthcoming Flora of Madeira. 

Hymenophyllum maderense Gibby & Lovis sp. nov. 

Habitu et textura H. wilsonii Hooker similis, praecipue venis segmentorum apice non attingentibus, 
cellulis laminariis brevioribus (ad duplo longioribus quam latioribus), minus regulariter dispositis, 
indusii ovatis, differt. Filix amphidiploidea, 2n = 62. (Owing to the omission of the expression 
"sp." in the Alphabetical List of Taxa with data given in Manton et al. 1986, the chromosome 
counts for H. maderense appear to be attributed therein to H. wilsonii). 

Holotypus: ex hort. Leeds, 1973, origin: Madeira, Queimadas, from a rock face in a gully below 
a levada, coll. J.D. Lovis s.n. 25 July 1970. BM (Fig. 1) 

Mat-forming perennial with slender branched rhizomes bearing thin translucent fronds. Fronds 
1.5 - 9.0cm in length, deep olive-green, oblong to oblanceolate in shape, pinnately divided, 
but sometimes with one or more lower pinnae much longer and more divided than the rest, 
giving a branched appearance; pinnae divided into 2-5 segments, segments broadest at the 
middle and about twice as long as broad; segmental vein terminates 4 cells short of end of 
pinna segment; lamina cells up to twice as long as broad, arranged haphazardly, each cell containing 
many (c. 80) chloroplasts, each chloroplast c. 5um in diameter. Sori appearing stalked, inserted 
perpendicularly to the plane of the lamina; indusial valves spreading apart at maturity, ovoid, 
not tapering markedly at the apex, margins entire. Spores c. 55^m in diameter (range 53-57). 
2n = 62. 

H. maderense is similar to H. wilsonii Hooker, but can be distinguished as the vein 
of the pinna segments stops short of the apex, the lamina cells are shorter and arranged 
haphazardly (as in H. tunbrigense (L.) Sm.), not in rows lying at an angle to the vein, 
and the indusial valves are ovate, not flask-shaped, and tapering markedly to the apex 
as in H. wilsonii. H. maderense is very rare in Madeira, being recorded only from a rock 
face and the side of a levada at Queimadas. However, the occurrence of hybrids of H. 
maderense x H. wilsonii (see below) indicates that it must once have been, and possibly 
still is, more widespread. Cytological evidence indicates very clearly that it is a markedly 
dibasic allopolyploid, derived by chromosome duplication from a hybrid of H. tunbrigense 
(2n = 26) x H. wilsonii (2n = 36). 

Hybrids of H. maderense *H. wilsonii have been discovered at Queimadas and at Pico 
do Tapeiro. These plants are very close to H. wilsonii in morphology, and can be distinguished 
only by their different chromosome number (2n = 49), an irregular meiosis and bad spores. 

Asplenium trichomanes subsp. maderense Gibby & Lovis subsp. nov. 

Habitu et textura A. trichomanes subsp. quadrivalens similis, praecipue frondis perviridibus 
subnitidis in statu vivo differt. Sporae 35 - 40um longae. Filix hexaploidea, 2n = 216. 



286 



FERN GAZETTE: VOLUME 13 PART 5 (1989) 




FIGURE 1. Hymenophyllum maderense, cult. Leeds 1973. A, whole frond x 1.3; B, portion of frond 
showing sori x 4. 




FIGURE 2. Asplenium trichomanes subsp. maderense, A & B wild collections; C ex cult. Leeds. 



NEW FERNS OF MADEIRA 



287 



Holotypus: Madeira, Pico do Ferreiro, in a horizontal rock crevice, c. 1400m alt., coll. J.D. 
Lovis s.n., 26 July 1970. BM (Fig. 2). 

Between A. anceps and A. trichomanes subsp. quadrivalens in morphology, but much closer 
to the latter taxon, from which it can usually be distinguished in fresh condition by darker 
green, sub-shiny pinnae. Spores 35 - 40^im long, rhizome scales < 4.5mm, narrowly lanceolate 
to subulate, with broad occluded central stripe. Hexaploid, 2n - 216, endemic to Madeira. 

Asplenium trichomanes subsp. maderense is believed to be an allopolyploid, derived 
from a hybrid of A. anceps and A. trichomanes subsp. qdbdrivalens by chromosome 
duplication. In addition to the type locality, it has also been discovered on Pico do Ariero 
c. 1800m alt. (coll. R. & W. Jager, 15 March 1981). 

Ceterach lolegnamense Gibby & Lovis sp. nov. 

Filix inter C. aureum (Cav.) L.v. Buch var. aureum et C aureum var. parvifolium Benl & 
Kunkel magnitudine intermedia, frondis plerumque 9 - 24cm longis, 1.5 - 3.8cm latis, sporis 

(33-) 35-42 (-45)um longis (perispore exclusus), 
numero chromosomatorum (2n = 216) 
magnitudine intermedius; distributi 
geographica differt. 

Holotypus: Madeira, Serra d' Agua, coll. J.D. 
Lovis s.n. 4 April 1969. BM (Fig. 3) 
The name lolegnamense is derived from an old 
name for Madeira (cf. Cossart 1984, pp. 2-4). 

Differs from Ceterach aureum (Cav.) L.v. Buch 
var. aureum and var. parvifolium Benl & 
Kunkel in size, with fronds usually from 9 - 
24cm long and 1.5 - 3.8cm wide, in spore size, 
(33-) 35 - 42 (-45)pm (excluding the perispore), 
chromosome number, 2n = 216, and 
geographical distribution, being endemic to 
Madeira. 

It has been suggested (see Manton et 
al. 1986) that C. lolegnamense is derived 
by hybridization between C. aureum var. 
aureum, a tetraploid taxon (2n = 144), 
and C. aureum var. parvifolium, an 
octoploid (2n = 288). C. lolegnamense is 
intermediate between these two taxa in 
size, spore size and chromosome number, 
but is confined to Madeira, whereas C. 
aureum var. aureum and C. aureum var 
parvifolium are known only from the 
Canary Islands. Cytogenetic studies are 
in progress to confirm the relationship of 
the three taxa, and it is intended that the 
two taxa from the Canary Islands will be 
recognised at specific rank (Reichstein & 
Vida, pers. comm.). 

Polystichum x maderense Johnson 

Holotype; Madeira, Rabacal, G.C. Joad 
s.n. February 1866 K (Fig. 4) 



FIGURE 3. Ceterach lolegnamense, part of holotype. Polystichum maderense Johnson 

(1866) was described from two or three 




288 



FERN GAZETTE: VOLUME 13 PART 5 (1989) 



fronds of a single specimen collected by G.C. Joad at Rabacal, at the head of the Ribeira 
da Janela; Johnson was of the opinion that this fern was 'exactly intermediate between 
P. falcinellum Presl and P. angulare PresP [= P. setiferum (Forssk.) Woynar] (Johnson 
1866). In April 1969 J.D. Lovis, in the company of T. Reichstein and P.G.J, de Joncheere, 
found a hybrid at Rabacal with morphology intermediate between P. falcinellum and P. 
setiferum, which he subsequently found to be pentaploid (2n = 205), thus confirming the 
proposed parentage of P. falcinellum (2n = 328) x P. setiferum (2n = 82). In June 1969 
M.L. Nilsson discovered a plant with similar morphology on Pico do Ariero, and that 
was determined as P.maderense by Hansen (1970). Unfortunately only a single sterile frond 
collected by Joad is at Kew (although in his description Johnson refers to the arrangement 
of the sori and shape of the indusia), and the Nilsson specimen at Copenhagen has shed 
all its spores. Therefore the hybrid nature of this fern can be confirmed only in the Lovis 
collection, but morphologically all these specimens are very similar and clearly represent 
the same taxon, the correct name for which must be P. x maderense Johnson. Like P. 
falcinellum, P. x maderense is a Madeiran endemic. 







?■ 



fofc 



it 




FIGURE 4. Polystichum x maderense, holotype. 



NEW FERNS OF MADEIRA 



289 



At the end of his account of Aspidium, Milde (1867) refers to P. maderense and suggests 
it may be the same as Aspidium falcinellum var. subbipinnata Moore. The varietal name 
var. subbipinnata appears not to have been published by Moore - Menezes (1914) refers 
to var. subpinnatum (sic!) Milde. Because the note by Milde follows his account for Aspidium 
frondosum, Romariz (1953) includes in the synonymy of P. maderense 'Aspidium frondosum 
var. subbipinnata Moore in Milde?; P. falcinellum var. subbipinnata Moore?'. However, 
a specimen at Kew from the herbarium of Thomas Moore labelled in Moore's handwriting 
1 Polystichum falcinellum var. bipinnatum, Hort. Veitch 1856, ex Madeira', is similar in 
morphology to P. x maderense. The following description is based on all the specimens 
studied: 

Fronds coriaceous, up to 80cm. Stipe c. 1/3 length of frond, rhizome and lower part of stipe 
covered in brown triangular scales with a dark central stripe, scales higher up stipe and on 
rachis more sparse, concolorous, lanceolate with tapering points. Lamina oblanceolate - lanceolate, 
pinnate, pinnatifid, but bipinnate at base of lower pinnae; pinnae lanceolate but with innermost 
apiscopic segment enlarged, pinna segments cut 1/4 to 1/2 width of the pinnae, not overlapping, 
slightly toothed, tapering to a distinct apical spine. Indusium > 1mm, peltate with central dark 
spot, margin irregularly toothed with a few multicellular hairs. Spores abortive. 2n = 205. 

Another Polystichum hybrid is known only from three herbarium sheets at Kew, and 
may well have existed as a single hybrid plant. One specimen from Camacha, Madeira 
was collected by Miss Ellen Taylor s.n. 19 December 1865, from 'a single plant growing 







FIGURE 5. Polystichum falcinellum x ?, Taylor s.n., 19 Dec. 1865. 



290 FERN GAZETTE: VOLUME 13 PART 5 (1989) 



in a bed of Nephrodium falcinellum ' (Herbarium of Revd R.T. Lowe) (Fig. 5). A further 
specimen was collected by Revd G.N. Burningham, and, as stated in a letter (22 September 
1875) from Burningham that is attached to the sheet, 'was growing amongst many plants 
of Aspidium falcinellum in Madeira, and which Mr Lowe when last in Madeira thought 
might prove a distinct species'; the third specimen has no collector's label, but may be 
a further Burningham specimen. 

This hybrid certainly has P. falcinellum as one parent as it has a coriaceous lamina, 
and similar indusia. The identity of the second parent is much more difficult; the only 
other Polystichum species on Madeira are P. setiferum (which seems unlikely as P. setiferum 
x P. falcinellum is the parentage of P. x maderense, and P x maderense differs in that 
it does not have rounded, crowded or overlapping pinnules) and the endemic P. drepanum 
(Sw.) C. Presl [it would be surprising if P. drepanum were to be the second parent for 
two reasons:- first, it is most unlikely that this very rare species which is now confined 
to a high valley in north-west Madeira was ever growing naturally in the region of Camacha, 
and secondly, since the combination of P. falcinellum and the diploid P. setiferum produces 
P. x maderense, it is not likely that the Camacha hybrid, with its generally similar form, 
could be the combination of P. falcinellum and P. drepanum, because the latter is a tetraploid 
species with a markedly triangular frond and very open pinnation]. The third possibility 
is that the second parent is an as yet unknown, probably diploid, very likely extinct species 
of Polystichum. Because of the rarity and uncertain origin of this hybrid we refrain from 
giving it a name, but it is described below: 

Polystichum falcinellum x ? 

Fronds coriaceous, up to 100cm long. Stipe c. 1/3 total frond length, green; rhizome and stipe 
base not seen; stipe sparsely covered and rachis more densely covered with brown lanceolate 
scales with tapering points. Lamina oblanceolate, up to 16cm wide, pinnate, pinnatisect; pinnae 
tapering, pinnules symmetrical about pinna axis except for innermost acroscopic pinnule which 
is always larger than its neighbour; pinnules up to 8mm long (innermost acroscopic pinnule 
up to 14mm), crowded with innermost pinnules overlapping, ovate, toothed and with apical 
spine. Indusia 1mm, peltate with small central dark spot, margin irregularly toothed. Spores 
abortive. 

Madeira, Camacha, a single plant growing in a bed of Nephrodium falcinellum, Miss Ellen 
Taylor s.n. 19 December 1865. Herbarium of Revd R.T. Lowe. K (Fig. 5). 

ACKNOWLEDGEMENTS 
We thank Dr N.K.B. Robson who assisted with the latin diagnoses, A.C. Jermy for advice, 
Prof. T. Reichstein and Prof. G. Vida for information regarding their studies on Ceterach 
in the Canary Islands, and Dr W. Jager for loan of his example of Asplenium trichomanes 
subsp. maderense collected on Pico do Ariero. 

REFERENCES 
COSSART, N. 1984. Madeira, the island vineyard. Christie's Wine Publication. London. 216 pp. 
JOHNSON, J.Y. 1866. Some account of a new species of fern (Polystichum maderense) recently discovered 

in the island- of Madeira. Ann. Mag. nat. Hist. Ill, 17, 287. 
HANSEN, A. 1970. Beitrage zur Flora der Inseln Madeira, Porto Santo und Ilhen Chao (Desertas). 

Bocagiana (Funchal) 25: 1-17. 
MANTON, I., LOVIS, J.D., VIDA, G. & GIBBY, M. 1986. Cytology of the fern flora of Madeira. 

Bull. Br. Mus. nat. Hist. (Bot.) 15: 123-161. 
MENEZES, C.A. de, 1914. Flora do Archipelago da Madeira. Junta Agricola da Madeira, Funchal 

212 pp. 
MILDE, J. 1867. Filices Europae et Atlantidis, Asiae Minoris et Sibiriae. Leipzig 31 1pp. 
ROM ARIZ, C. 1953. Flora da Ilha da Madeira. - Pteridofitos. Revta Fac. Cienc. Univ. Lisb. IIC, 

3:53-112. 



FERN GAZ.. 13(5) 1989 291 



THE ECOLOGY AND DISTRIBUTION OF PTERIDOPHYTES OF 

ZOMBA MT., MALAWI* 

AUDREY BERRIE 
Biology Dept., Chancellor College, P.O. Box 280, Zomba, Malawi, Central Africaf 

ABSTRACT 

Ninety-six species of pteridophytes are known from the Zomba area, from the earliest 
collections of the Livingstone Expedition, 1859, up to recent collections by the author 
from 1980-86. Distributional data is given for the pteridophytes of Zomba Plateau (altitude 
1500-2085m) with species in remnants of undifferentiated Afromontane forest and 
grassland, riverine forest and plantations of introduced conifers, and the pteridophytes 
of the mountain slopes (1000-1500m) from the rocky cliff-edges, the Brachystegia/mixed 
woodlands and the plantations of the lower slopes. Very few pteridophyte species are 
found below 900m altitude in the surrounding savanna areas. Some species from earlier 
collections, e.g. 1896 (Whyte), 1967 (Berrie), have not been seen recently from 1980- 
86 and may no longer be present in the Zomba area. 

INTRODUCTION 
The earliest European exploration of Central Africa by the Livingstone Expedition passed 
through the Zomba area in September 1859 (Baker 1959, Jackson 1959). The plants collected 
then included a small number of pteridophytes (12 species in 10 genera) among the very 
many numbers of other plant groups (Burkill 1897, 1898 & 1906). Later collectors in special 
expeditions also only included a few pteridophytes e.g. the Brass collections of the Vernay 
Nyasaland Expedition of 1946, with 33 pteridophyte numbers (27 species in 17 genera) 
out of the 288 numbers of all plant groups from the Zomba area (Ballard 1953, Brass 
1953). Schelpe (1970) in Flora Zambesiaca listed only 21 species, in 14 genera, for this 
area. The most recent checklist of the pteridophytes of Zomba Mt. (Berrie 1984) included 
93 species in 43 genera. 

The present paper discusses the distribution of 95 pteridophyte species in 45 genera 
presently or previously recorded for the Zomba area. 

STUDY AREA 
Location 

Zomba mountain lies east of the Great Rift Valley of Central Africa, in the Southern 
Region of Malawi. At latitude 15°10' - 23'S and longitude 35°15' - 23'E it is the highest 
part and the most northern inselberg of the Shire Highlands. This ridge of highlands runs 
NE - SW between Lake Chilwa to the east and the Shire River to the west (Figure la). 

The Zomba mountain massif consists of two plateau areas separated by the Domasi 
River, flowing almost due east between the northern Malosa Plateau and the Southern 
Zomba Plateau (Figure lb). Both plateaux are forest reserves. Zomba Plateau is 8km N- 
S and 12km E-W, with a narrow road on the southern scarp from Zomba town at its 
base. There are many small roads around and through the conifer plantations on Zomba 
Plateau, interconnected by narrow footpaths. Malosa Plateau is larger, 14km N-S and 10km 
E-W, with no road up its very steep scarp, no regularly used pathways over or around 
it, and only a few trial plots of exotic conifers planted there. Due to its inaccessible nature 
and the limitation on time available for fieldwork the Malosa Plateau was not included 
in the present study. Reference to Zomba Mt. in the text thus includes Zomba Plateau, 
the south-west and south-eastern slopes, Zomba town area at the southern base of the 
mountain, and Zomba District the area around the mountain massif. 

* First submitted September 1987. 

t Present address: Biology Dept., National University, P.O. Roma 180, Lesotho, Southern 

Africa. 



292 



FERN GAZETTE: VOLUME 13 PART 5 (1989) 




J3 



Us 



\i~ 










(0 



General vegetation 

Malawi is within the floristic region or phytochorion known as the Zambesian Region 
in High Africa (White 1983). On the "Vegetation Map of Africa" (UNESCO/AETFAT/ 
UNSO, White 1983) Zomba mountain is shown as a tiny island of 'undifferentiated montane 
vegetation, (a) Afromontane', No. 19 in the series Forest Transitions and Mosaics (No. 
11-24). This small area, 19a, is surrounded by 'wetter Zambesian miombo woodland 
(dominated by Brachystegia, Julbernaldia, and Isoberlinia)', No. 25 in the Woodland Series 
(No. 25-30). 



PTERIDOPHYTES OF ZOMBA MOUNTAIN 



293 



Zomba Plateau has a number of vegetation types including small areas of montane 
grassland, a few isolated patches of montane forest, and riverine forest along the main 
Mulunguzi River valley and a few streams. The largest areas of conifer plantations are 
Pinus patula Schlecht. & Cham, with other older trial plots of exotics, mainly conifers, 
and a few of the indigenous conifer Widdringtonia nodiflora (L.) Powrie (Mulanje Cedar). 

The Afromontane vegetation of Zomba Plateau includes grassland along the western 
rocky ridges from about 1800-2085m. The rough grassland with small shrubs and scattered 
herbs has species of Protea, Xerophyta (Vellozia), Vernonia, Senecio, Helichrysum, and 




FIGURE 2. Diagrammatic representation of Zomba Mt. showing the altitudinal areas of Zomba Plateau 
(ZP). Zoimba slopes (ZS). Zomba town (ZT). and Zomba District (ZD). (Abbreviations: Peaks 
on Zomba Plateau: R = Radiomast and Malumbe Peak (2085m): C = Chiradzulu Peak (2050m): 
F = Nawimbe Firetower (1800m): M = Mulunguzi Peak (1769m). Lookouts: Q = Queen's View; 
E = Emperor's View. TP = Trout Ponds. CHF = Chingwe's Hole Forest. CV = Chivunde 
valley. In Zomba town. FR = Forestry Research Institute of Malawi (F.R.I.M.): M = The National 
Herbarium of Malawi (MAL). CC = Chancellor College. University of Malawi. 



294 FERN GAZETTE: VOLUME 13 PART 5 (1989) 



Plectranthus. The tussocky grasses are dominated by species of Andropogon and Exotheca, 
while the forest/grassland marginal areas have tall plants of Hypericum sp. and shrubs 
of Philippia benguelensis with Bracken, Pteridium aquilinum, dominant in this transitional 
zone. The northern and eastern rims of the high plateau are almost completely planted 
with Pinus patula, except for remnants of riverine forest in strips along the smaller Mulunguzi 
tributaries. A very narrow fringing plantation of Eucalpytus sp. is along the southern rim, 
almost at the cliff-edge, around Mulunguzi peak (1769m) near the Lookout area, about 
750m above Zomba town (Figure 2). 

The remnants of Afromontane forest on Zomba Plateau are found at the heads of 
small valleys, hanging at the cliff-edges, with small streams falling to the main valleys below, 
as at Chingwe's Hole Forest (1900 alt.) facing NNW. The stream here forms the headwaters 
of the Namitembo River, flowing westwards to the Shire River. Similar remnant forest 
is found where the Chivunde stream begins, flowing due north to join the Domasi River 
in the valley below. 

Chingwe's Hole (montane) Forest, Figure 3, includes very many tree species evenly 
spread throughout a very small area with phorophytes ('host trees') for epiphytic ferns 
including Podocarpus latifolius, Myrica salici folia, Nuxia congesta, Cassipourea congoensis, 
Diospyros whyteana, Ilex mitis and Xymalos monospora. A few genera of trees found 
here (Halleria, Ilex and Nuxia), have the same species along the river but one genus, Syzygium, 
has one species in the montane forest (S. guineense subsp. afromontanum) and another 
(S. cordatum) in the river forest. Halleria lucida is found usually at the montane forest 
margins and occasionally in small clumps on the nearby grassland, but along the river 
it is very abundant and bears the greatest number of epiphytic ferns, of most species, in 
the riverine forest. 

Riverine forest occurs along the Mulunguzi River and its smaller tributaries on Zomba 
Plateau, and extends downriver from the Mulunguzi Dam at the plateau rim, about 1450m 
alt., into the deeply cut valley below as far as the top of the gorge, at approximately 1380m. 
Even on the Plateau the riverine forest is mostly a fringing forest. There are a few dense 
forest patches where Ilex mitis and Rauvolfia caffra bear filmy ferns and other epiphytes, 
but along the river banks there are smaller trees, of Halleria lucida and Syzygium cordatum 
bearing many epiphytes on the trunks and branches, but not the filmy fern masses of the 
closed forest. 

The very steep slopes above the Gorge and the exposed rocky edges of the Plateau 
have a rather open woodland of Brachystegia/ mixed species. Only a few epiphytic ferns 
occur here on trees of Julbernaldia globiflora, Hetermorpha arborescens, Parinari 
curatellifolia and Brachystegia spp. The very occasional trees of, Fig. 4, Acacia sieberana 
scattered on the upper mountain slopes carry a very heavy load of fern epiphytes. The 
cliff-faces on the southern scarp stand out above the very stunted woodland at their lower 
edges, about 1300m alt. The same tree species are here very small, sometimes only 2-3m 
in height, and rarely with any epiphytic ferns though they sometimes have an occasional 
orchid present. The conifer plantations, Fig. 5, of the lower slopes are mostly Pinus patula, 
with a small amount of Cupressus sp., stretching from the town boundary at 1000m up 
to the stunted woodland at the cliff-base. Below the 1000m level, within and around Zomba 
town, the land is highly cultivated with Maize and other crops except along the streams 
ana the Mulunguzi River. Remnants of riverine forest include some large forest trees like 
Khaya nyasica and Filicium decipiens even in the town area. Away from permanent streams 
there are not many places where pteridophytes can survive. Some of the very dry rocky 
outcrops on the lower slopes of the mountain and at some distance from its base have 
poikilohydrous, xerophytic ferns only seen easily during the wet season. The open savanna 
areas surrounding the mountain have very little uncultivated land. Only streamsides and 
water channels support any vegetation in which pteridophytes may be found. The Mulunguzi 



PTERIDOPHYTES OF ZOMBA MOUNTAIN 



295 




FIGURE 3. Montane epiphytes in Chingwe's Hole Forest (altitude 1900m) on Zomba Plateau. On 
Myrica salicifolia, V/ 2 - 2m from ground level, there are small narrow pendant fronds of Asplenium 
sardersonii (centre and lower branch) with proliferating frond tips, and the larger spreading 
fronds of Arthropteris monocarpa projecting from long creeping rhizomes closely attached to 
the tree trunk, (photo: G.K. Berrie) 



296 



FERN GAZETTE: VOLUME 13 PART 5 (1989) 




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PTERIDOPHYTES OF ZOMBA MOUNTAIN 297 



R. is permanent but many of the smaller streams dry up completely during some part 
of the year. Thus the vegetation of Zomba Mt. while going through severe conditions during 
the dry season appears always green, but below the 1000m level, months of dry season 
change the appearance of the land around the mountain, each year. 

Climate 

The climate of Malawi is the Tropical Summer Rain, type II, of Walter (1971), with the 
wet summer season of the southern hemisphere from November to April and the dry season 
from May to October. 

However, the dry season can be divided into the cool, early dry season months and 
the later, hot to very hot, months. In Zomba the coldest months are usually June/July 
(minimum 6°C) while the hottest months are Sept./October (maximum 34°C). Frosts have 
not been recorded for Zomba town, or at Chancellor College (CC) where meteorological 
records have been taken since 1972, but frost damage to tree ferns (Cyathea sp.) has been 
seen on Zomba Plateau (ZP), at 1500m and 1800m, on a number of occasions during 
late June/early July in recent years between 1980-86. 

Continuous temperature records are not available for Zomba Plateau but some data 
(F.R.I.M.) for 1978-80 can be compared with that for Chancellor College (CC: Geog. 
Dept.), as follows (in °C): 



Abs. 


Abs. 






Abs. 


Abs. 




max. 


min. 


Range 




max. 


min. 


Range 


28 


4 


24 


wet season: 


27 


11 


16 


34 


6 


28 


wet season: 


33 


11 


22 



ZP. (1900m) dry season: 
CC. (900m) dry season: 

Rainfall data for both places for the ten year period 1976-86 gives a mean annual total 
for ZP as 2244mm and for CC as 1464mm. If a dry month is one with less than 25mm 
then the dry season may be given as 5 months (May - Sept.) or 6 months (May - Oct.) 
since October rainfall varies from - 132mm on ZP and from - 72mm at CC in different 
years. 

An additional factor in climate is the frequent mist and cloud-cover on the mountains. 
The cloud base may extend down to the 1000m level at the base of the mountain, around 
the edge of Zomba Town. During some wet season months (Jan. - March) there may 
be 10-13 complete days like this; in dry season months (June - Sept.) there may be 6- 
8 days per month like this, or for most of the morning only. Looking up to Zomba Mt. 
from Chancellor College in the dry season (Aug.) the very early morning sky (5.30-6.30 
am) may be clear all around the mountain, but low heavy clouds come across from the 
eastern end, from over Lake Chilwa, so by 7 am the Plateau is enveloped in thick cloud. 
This may remain to 10 am or midday. This 'condensation cloud' must add considerably 
to available water, especially for the epiphytes on the Plateau and upper slopes, at a time 
when measured rainfall is zero or nearly so. Cloud cover can also change very rapidly 
during the wet season, giving heavy isolated local rain, e.g. on the lower slopes near the 
Herbarium but none in Zomba town centre about 1km away, or at Chancellor College 
1.5km lower down the slope. 

This pattern means that the climate controls the seasons but the local weather conditions 
control the rainfall, so the weather as much as the climate affects the plant distribution. 



298 



FERN GAZETTE: VOLUME 13 PART 5 (1989) 




PTERIDOPHYTES OF ZOMBA MOUNTAIN 



299 



OBSERVATIONS ON THE DISTRIBUTION OF THE PTERIDOPHYTES 
Habitat 

Most pteridophyte species are not restricted to one vegetation type, i.e. to montane grassland, 
montane forest, riverine forest, woodland or the plantations described. Instead they occur 
in similar habitats within and around these areas. Here I have grouped the pteridophytes 
by habitat, from wet to dry; from the permanently wet, closed forest type (montane), the 
permanently moist, closed forest (riverine), to the remnant riverine forest, and the woodland 
habitats. These are listed as Forest/woodland species in Habitats I - VI (Table 1). Other 
permanently wet, non-forest habitats include riversides, marshlands and swamps (Habitat 
VII). The seasonally very wet habitats are VIII: seasonal pools, and IX: seasonal flush 
areas on rocky outcrops. The remaining habitats are not particularly wet and are without 
closed tree-cover, but may be found at edges of various forest types and in more open 
places. These are X: marginal habitats, XI: sheltered habitats, and XII; exposed habitats. 

Altitude 

Table 1 lists the distribution of the pteridophytes in these Habitats, 1-XII, in each altitudinal 
area, with details of habit and habitat of each species. The whole area was divided into 
four altitudinal zones as follows: 

1. Zomba Plateau (ZP), all above 1500m alt. up to 2085m; and the area around the Mulunguzi 
Dam, at approximately 1450- 1500m; 

2. Zomba slopes (ZS), the mountain slopes from 1000- 1500m; 

3. Zomba town (ZT), the inner town area, approximately 900-1000m; 

4. Zomba District (ZD), the lower slopes and surrounding area within the administrative 
boundary of Zomba District, down to 650m alt. 

Some species are restricted to each of these four areas, to ZP, ZS, ZT, or ZD. Others 
have wider distribution, so combinations of these areas give continuous distribution of species 
in ZP,ZS,ZT,ZD; ZP,ZS,ZT; ZP,ZS; ZS,ZT,ZD; ZS,ZT; and ZT,ZD; while discontinuous 
distribution occurs in ZP,ZS,ZD; ZP,ZT,ZD; and ZP,ZT. 

This altitude zonation is used in the distribution diagrams showing combinations of 
Habitats and altitude in Figure 6. 



ZP 



ZS 



ZT 



ZD 



Habitat 


3 


I 


-VI 


» 




























forest/woodland 




Separate Habitats I - VI 


species 








I 


II 


III 


IV 


V 


VI 




f 




^ 


















r \ 










22 


4 




6 




5 


3 


4 


1 




1 


9 


l 




4 




4 




4 




6 










1 








l 




4 




4 
















































4 




1 












l 


4 






















1 


1 










1 


l 




22 +1 + 4 +6 +1 


5 


3 


4 +1 


I 


S +1 +L +1 


4 +4 


=» 34 spp. 






= 5 




= 12 


- 8 














5 


+ 3 


+ 5 








t-i; 


> 






+8 







= tot;:! 3-1 app , 



300 



FERN GAZETTE: VOLUME 13 PART 5 (1989) 



b Habitats I - VI, forest/woodland species 







I 


II 


III 


IV 


\ 


' 


: VI 






ZP 




5M 


3KRC 


4-MR 


1M+R 


1R+C 


9R 


1R 


4 M+ R+Rr+W 




4M+R+Rr+W 






























zs 










1R 






1R 


4 R+Rr+W 




4R+Rx+W 






















ZT 














1H 


4 R+Rr+W 






















ZD 












1R 


1R 








5 


+ 3 


+ 5 


+ 1 


+ 12 


+8 m 34 total. 


M - 5 


+3 


+ 5 




2 


+4 +4 = 21; 

I spp. in H»montane forest; 


R - 


+3 


+5 


+1 

29 


+12 +4 +4 = 29; 

spn. in R, riverine forest; 


G » 


+ 3 





+ 1 

4 


0=4; 
spp. in 0,nlif f-ed're forest; 


Rr = 








+4 +4=8; 
8 spp. in Rr, rerun-Tit riverine forest; 


W - 










8 spp 


+4 +4 = 8 
. in W, wood lands. 


; 



32 



10 



= 2 



c 


Habitat VII jriversic 


es 






p 


oo Is 
VIII 


Habitat 


IX:fl 


ush 


ZP 

- 11 


1 


2 




1 




1 




r 1 
5 




1 






1 


1 
































zs 

- 16 


3 


2 




1 






5 






2 




3 








1 




1 




2 












i ; 




























ZT 
- 10 


.1 


2 




1 




1 




2 




3 






2 


1 




1 




2 




























ZD 
- 4 




I 




1 




2 




2 




1 




1 






1 +3 +1 
totals: = c 


*2 

>0 


si 


fl 

>P 




+1 




+ 5 




+ 1 




+ 2 




+ ;^ 




2 
_2_ 


1 +2 
- 7 


+ 1 
sp 


). 


+1 




+ 


2 



= 4 



PTERIDOPHYTES OF ZOMBA MOUNTAIN 



301 



ZP 



zs 



ZT 



ZD 





Habita 


t 


X: 


marg 


inal 


Ha 


bi 


ta- 


t : 


T 




shelte 


red 


Habitat 


XIT: 


s> 


po 


> 

13 


2 


2 




4 




5 




1 




2 




1 


-4 


1 


7^ 

2 




4 












































X2 


1 


2 




4 




5, 




1 




2 




1 




1 




2 


- 7 




2 




4 










































6 




2 




4 




4 


1 




2 




1 




2 


=10 


1 


2 


























l 
2 




2 






1 




1 


= 2 


1 


to 


2 +1 
tals:» 




>2 


3 P1 


h4 

3. 


H 


^5 


4 


► 1 


= 11 


+ 1 

SPD 


+ 1 


+ 2 




1 +1 +1 +2 

= 9 SDp 


•♦ 


4 





= 6 



= 1 



ZP 



ZS 



ZT 



ZD 



26(+lv) 



All Habitats 
7^S 



7(+lv) 



4 4 



F^-S — P*"*T 



14 



14 



14 



21 



2 ^ 



= 68 spp. (72%) 



i 1 



1 



S2 



= 55 spp. (58%) 



39 spp. (41%) 



1 =15 spp. (16%) 



26 +4 +7 +4 +4 +14 



21 



1 



1 = 95 spp 



FIGURE 6. Distribution of pteridophyte species in Habitats I-Xii in altitudinal zones (ZP, ZS, ZT, 
ZD) showing restricted, continuous and discontinuous patterns: a) forest/woodland species 
(Habitat I-VI); b) same in different forest types; c) non-forest, permanently wet, riversides (Habitat 
VII); seasonal pools (VIII); and seasonal flush (IX); d) marginal (X); and seasonally extreme, 
sheltered (XI) and exposed (XII); e) species in all habitats. 



302 



FERN GAZETTE: VOLUME 13 PART 5 (1989) 



TABLE 1 



Distribution of the pteridophytes by habitat and altitude. The altitudinal areas in each Habitat are 
separated by a dotted line; the Habitats are separated from each other by a solid line. See text for 
abbreviations. Taxa not seen during this study (1980-1986) are marked with an asterisk* 

Nomenclature follows Flora Zambesiaca (Schelpe 1970) except for the family Thelypteridaceae which 
follows Holttum (1974) in the recognition of more genera. 

(3) 



Plant habit 
and habitat 



(1) 


(2) 


Habitats 


Local distribution 


I-XII 


habit 


Area, and 


and 


species 


Regional distrib. 



I: montane forest only (ZP only: 5 spp. including 1 var.) 
ZP only = 5 spp. 



Asplenium boltonii 


f:T/E 
Za.Zb. 


A. dregeanum 


r:T/E 
Za.Zb.M.SA 


Elaphoglossum 
acrostichoides 


r:E 
Zb.L.M.SA. 


A. lobatum 


r:T 
Zb.M.S.SA. 


Dryopteris 

inaequalis 

var. atropaleacea 

(see Habitat X for other 


r:T 
Za. 


var.) 





Terrestrial in deep shade or low level epiphyte, to lm 
on tree trunks. 

Terrestrial, or epiphyte on tree bases. 



Epiphyte, on upper surface of branches, 
2-3m from ground level. 

Terrestrial, in deep shade near stream. One 
location only 

Large erect terrestrial, like a small tree fern, 
in deep shade; few scattered plants, one 
location only. 



II : montane and riverine forest, and cliff-edge forest (ZP only) 
ZP only = 3 spp. 



Elaphoglossum 
spathulatum 

Hymenophyllum 
polyanthos 
var. kuhnii 

Oleandra 
distenta 



r : E/L Epiphyte in montane and cliff-edge forest; 

Za.Zb. L.M.S.SA. lithophyte in riverine forest. 

a : E/L Pendant epiphyte, massed on tree trunks up to 

Zb.M.SA. 3m, on vertical and horizontal surfaces; 

lithophyte near waterfalls; usually in deep shade. 

o to a : E/T/L Epiphyte, becoming terrestrial from rooting of 

Za. Zb.M.S.SA. pendant rhizomes. Old erect rhizomes covered by 

lichens growing at cliff-edge; lithophyte at riverside. 



Ill : montane and riverine forest (ZP and ZS = 5 spp.) 
ZP only = 4 spp. 



Asplenium 


a:E 


mannu 


Za.Zb.M. 


A. megalura 


o-:E 




Za. 


A. sandersonii 


a:E 




Zb.M.SA 


Trichomanes 


o: E 


borbonicum 


Zb.M.SA 



Small creeping epiphyte; ground level to 4m, 
on large and small trees and lianes; often with A. 
sandersonii. 

Tufted epiphytes; small fronds (10-15 cm) at 

l-3m from ground but much larger (to 40cm) at 6m 

and in higher branches. 

Spreading, low level epiphyte; l-2m from ground. 

Tiny semi-erect filmy fern, low level epiphyte 
l-2m from ground; usually with Hymenophyllum 
polyanthos. 



PTERIDOPHYTES OF ZOMBA MOUNTAIN 



303 



III : montane forest (ZP) and riverine forest (ZP, ZS). 
ZP, ZS = 1 sp. 



Belchnum 
giganteum 



f : T Large erect terrestrial; on open riverbanks 

Za.Zb.L.M.S.SA and deeply shaded streams. 



IV : riverine and cliff-edge forest (ZP only) 
ZP only = 1 sp 



Blechnum 
tabulare 



o:T 
Za.Zb.L.M.S.SA. 



Erect terrestrial; in forest/woodland at cliff-edges 
and along streams. 



V : riverine forest only (ZP, ZS, ZT and ZD 
ZP only = 9 spp. 

Blechnum attenuation f to a : E 

Za.Zb. 



12 spp.) 



Lycopodium 
verticillatum 


o:E 
Za.Zb.L.M.SA, 


Blotiella 


r:T 


natalensis 


Za.Zb.M.SA. 


Elaphoglossum 
hybridum 


r.T 
Zb.M.SA. 


*Asplenium erectum 


s* 




Za.Zb.M.S.SA. 


* A. linckii 


s* 




Zb. 


*A. monanthes 


s* 




Zb.L.M.SA. 


*A. pseudoauriculatum 


s* 
Zb.M.SA. 


* Polypodium 
polypodioides 


s* 
Zb.M.S.SA. 


subsp. ecklonii 




V : as above 




ZD : 1 spp 




*Pteris vittata 


s* 



Erect epiphyte on tree ferns (Cyathea spp.),, 
ground level up to 2.5m. 

Epiphyte, on horizontal branches and tree 
trunks at angles up to 70° but not on vertical trunks; 
young plants erect, later bending; mature plants 
pendant. 

Large terrestrial, like a small tree fern, 
in deep shade, swampy area in wet season. 

Tufted terrestrial; on vertical earthbank 
at riverside, in deep shade. 

21-7-62, Mwanza 2512 (MAL*), 
"on rocky riverbank". 

1896, Whyte (K*) "Zomba Rock" 



1-4-80, Blackmore 1166 (MAL*), 
"streamside in dense forest" 

28-5-46, Brass 16059 (K*, NY), 
"occasional on rocks in riverine forest" 

21-1-67, Berrie 30 (K*), 

"epiphyte on tree branch over-hanging river' 



5-4-80, Blackmore 1193 (K, MAL*), 
Za.Zb. L.M.N. S.SA. "by roadside in riverine forest dominated by Khaya 
nyasica". 



V : as above 
ZP, ZS = 1 sp. 

Selaginella 
kraussiana 



a : T Creeping terrestrial, along riverbanks and 

Zb.M.S.SA. masses in ground flora of forest. 



304 



FERN GAZETTE: VOLUME 13 PART 5 (1989) 



V : as above 
ZT, ZD = 1 sp 

*Asplenium formosum 



Za.Zb.M. 



(ZT):8-7-62, Mwanza (MAL*), "rocky surface near 

River" 

(ZD): 11-4-80, Blackmore and Morris 1223 

(MAL*), "riverbank in Khaya forest" 



VI : montane forest (ZP), riverine forest (ZP, ZS), remnant riverine forest (ZP, ZS« ZT), and woodlands 
(ZS). (all areas = 8 spp!) 
ZP, ZS, ZT = 4 spp. 



Asplenium theciferum 
var. concinnum 


a:E 
Za.Zb.M.SA. 


Pleopeltis 
macrocarpa 


ftoa:E/L 

Za.Zb.L.M.S.SA. 


Loxogramme 
lanceolata 


f:E/L 
Za.Zb.M.SA. 


Asplenium 
aethiopicum 


f : E/T/L 
Za.Zb.L.M.S.SA. 



Epiphyte, low and high levels, trunks and 
branches; in shade of forest, in open and closed 
woodland, occasionally on road-side trees (ZT). 

Epiphyte, usually on upper surface and 
sides of branches, occasionally on tree trunks; 
lithophyte near river at lower altitudes (ZS, ZT). 

Low level epiphyte in deeply shaded montane 

forest (ZP), and near river (ZP); lithophyte by river at 

lower altitudes (ZS, ZT). 

Epiphyte on mid to high branches from 4- 10m, 
in forest (ZP, ZS) and at riverside (ZP, ZS); on tree 
ferns (Cyathea, ZP), and cycad trunks 
(Encephalartos, ZT). T/L on ZS, ZT. 



VI : as above 
ZP, ZS = 4 spp. 




Pleopeltis 
excavata 


a:E/L 
Za.Zb.M.SA. 


Pyrrosia 
rhodesiana 


o:E/L 
Zb.M. 


Arthropteris 
monocarpa 


a:T/E 
Za.Zb.M.SA. 


Asplenium 
anisophyllum 


f:T 

Zb.M.S.SA. 



Epiphyte, on medium to high level branches on 
ZP, lower levels in woodlands (ZS); rarely lithophyte 
at lower altitudes (ZS). 

High level epiphyte in montane forest (ZP), 

lower level epiphyte or lithophyte at lower altitudes, 

near rivers. 

Terrestrial, becoming epiphytic; in shade of 
forest (ZP) and in open woodland (ZS), and outer 
parts of conifer plantations (ZP, ZS). 

Terrestrial, tufted but spreading; in general 
ground flora of montane forest (ZP), on steep banks 
of riverine forest (ZP), on rocky slopes of closed 
woodland (ZS). 



VII : riversides, streambanks, marshlands and swamps (all areas = 20 spp.) 
ZP only =1 sp. 



Lycopodium a : T 

carolinianum Za.Zb.M. 

(var. affine/tuberosum?) 



Creeping terrestrial; montane grassland 
streamside, very marshy, with Sphagnum sp. 
(bog moss). Some plants with tubers. 



VII : as above 
ZS only = 3 spp. 

Vittaha volkensii 



r(s):E 
Zb.M. 



Mid-level epiphyte, 3-4m from ground, on trunk 
and large branch (crown base) of Parinari tree near 
waterfall. First collection for Zomba: 28-5-85, Berrie 
636 (MAL). 



PTERIDOPHYTES OF ZOMBA MOUNTAIN 



305 



Cheilanthes 
farinosa 



*Pteris dentata 



r(s):T 
Za.Zb.B. 



Za.Zb.L.M.SA. 



Small terrestrial, in rock crevice at edge of 
woodland, 3m from river. First collection for Zomba: 
28-5-85, Berrie 637 (MAL). 

31-12-81, Chapman & Patel 6029 
(MAL*), "along strearnside, close to water" 



VII : as above 
ZT only = 1 sp 

Equisetum 
ramosissimum 



f to a : T Terrestrial with spreading, deep underground 

Za.Zb.L.M.SA. rhizomes; along streambanks of lower mountain 

slopes; always in moist shady positions. 



VII : as above 

ZP, ZS, ZT = 2 spp. 

Cyathea dregei 



Tectaria 
gemmifera 



a to r: T/A 
Za.Zb.L.M.S.SA. 



ftoa:T 

Za.Zb.M.SA. 



Tree fern; terrestrial, but may stand in water at 
riverside; always very close to water; few by montane 
grassland stream, abundant by river (ZP); not in 
montane forest; rare on lower mountain slopes, one 
by river, ZT. 

Large tufted terrestrial with very spreading 

fronds; along river-side paths (ZP), streambanks (ZS, 

ZT); always in moist shady places. 



VII : as above 
ZP, ZS, ZD = 1 sp. 

Osmunda regalis 



a : T/L/A/E Usually terrestrial by river (ZP), semi-aquatic 

Za.Zb.L.M.S.SA. between rocks, lower parts submerged in wet season; 

in slow moving water, not in rocky rapids; 

occasionally epiphytic on riverside tree ferns 

(Cyathea, ZP). 



VII : as above 
ZP, ZT, ZD = 1 sp. 

Cyclosorus 
interruptus 



a : T/A Creeping terrestrial; underwater in wet season; 

Za.Zb.B. M.N. SA. riverside swamp (ZP), marshy stream on lower slopes 
(ZT), silted streamside (ZD). 



VII : as above 
ZP, ZS, = 5 spp. 




Cyathea thomsonii 


o:T 
Za.Zb.M. 


Pneumatoptehs 
unita 


r :T 
Za.Zb.M.SA 


Diplazium 
zanzibaricum 


r:T/A 
Za.Zb.M.SA. 


Marattia fraxinea 


a: T/A 
Za.Zb.M.SA. 



Tree fern; terrestrial, usually in shade at 
water's edge. 

Terrestrial; on extreme edge of riverbank, 
extending over the water. 

Terrestrial, like a small tree fern; in deep 
shade by river (ZP); by stream between conifer 
plantation on mountain slopes, just in water in wet 
season. 

Terrestrial, extremely large, spreading; 

almost in water at riverside, usually never more than 

5m from water's edge. 



306 



FERN GAZETTE: VOLUME 13 PART 5 (1989) 



Thelyptgeris 
confluens 



o to a : T/A Terrestrial to semi-aquatic; in river source 

Za.Zb.B.L.M.N.S.SA. (Mulunguzi Marsh), edges of Dams (ZP); at river- 
side (ZS), swampy in wet season. In slight shade or 
exposed positions. 



VII : as above 
ZP, ZT = 1 sp 

Amauropelta 
bergiana 



r:T 
Za.Zb.M.S.SA. 



Terrestrial at riverside (ZP), by stream (ZT) 



VII : as above 

ZS, ZT, ZD = 2 spp 

Christella 
dentata 

Pteris friesii 



f : T Terrestrial at streamside; in slight shade or 

Za.Zb.B.M.SA. sheltered positions. 

o to f : T Terrestrial; in deep shade in riverine 

Za.Zb.M.SA. areas, at bases of rocks, at edge of old Cupressus 

plantation. 



VII : as above 
ZS, ZT, = 3 spp. 




Asplenium 
pumilum 




o:T 
Za.Zb. 


Microlepia 
speluncae 




r:T 
Za.Zb.B.M.N.SA 


Pellaea 
doniana 




f:T 

Za.Zb.M. 



Usually terrestrial, in deep shade; on moist 
earthbank, at riverside; by small stream; on 
stonework of river bridge, in moss layer. 

Terrestrial; rocky area near river, in sheltered 
positions at base of large rock. 

Terrestrial, in deep shade on riverside bank; 
occasionally in shelter of rocks on woodland slopes. 



VIII : seasonal pools (ZD only) 
ZD only = 2 spp. 



* Ceratopteris 
thalictroides 

*Marsilea sp' 



s* 10-8-71, Howard- Williams 229 (MAL*). 

Za.Zb.B.M.S.SA. "growing in mud . . . partly submerged" 

s* 3-2-55. Jackson 1453A (MAL*), 

Za.Zb.B.M.N.SA. "growing in water of rice gardens" 



IX : seasonal flush (all areas = 7 spp.) 
ZP only = 1 sp. 



Ophioglossum 
polyphyllum 


r:T 
Zb.B.M.N.SA. 


IX : as above 
ZT only = 2 spp. 




Actiniopteris 
radiata 


r:T 
Za.Zb.B.N.S.SA 


Isoetes 
abyssinica 


r:T/A 



Terrestrial, on rocky outcrop; at margin 
of montane grassland and montane forest. 



Terrestrial; rocky area with slight shade. 



Terrestrial; on level top of rocky outcrop; 
in shallow soil; water-logged in wet season. 



PTERIDOPHYTES OF ZOMBA MOUNTAIN 



307 



IX : as above 
ZP, ZS, ZT, ZD 

Ophioglossum 
gomezianum 



1 sp. 



a:T 
Za.Zb. 



Terrestrial; in shallow soil on slopes and tops 
of rocky out-crops; may be water-logged in wet 
season. 



IX : as above 

ZS, ZT, ZD = 1 sp. 

Actiniopteris 
dimorpha 



a : T Terrestrial; in outcrop areas, at edges of big 

Za.Zb. B.M.N.SA. boulders and in rock crevices; on rocky slopes below 
flush areas and on rocky roadside banks. 



IX : as above 
ZS, ZT = 2 spp. 

Ophioglossum 
costatum 



Selaginella 
njam-njamensis 



f:T/A 
Za.Zb. 



a:T/L 
Za. 



Terrestrial; on flat top of rocky outcrop, in 

shallow soil layer; usually water-logged in wet season; 

with Isoetes sp. 

Terrestrial; edges of flush area; in small pockets 
of soil on rocky outcrop, extending growth over 
surrounding bare rock surface. 



X : marginal habitats (all areas = 14 spp. including 1 var.) 
ZP only = 2 spp. 



Athyrium 
schimperi 



Cheilanthes 
inaequalis 



o:T 
Za.Zb.L.SA. 



o:T 
Za.Zb.B.M.SA. 



Terrestrial; edges of forest and grassland 
and at cliff— top, often large plants; occasionally at 
riverside similar; very small plants under rock ledges 
on montane grassland and on roadside banks (ZP). 

Terrestrial; forest margins and edges of conifer 
plantations; in partly shaded positions. 



X : as above 
ZS only = 1 sp. 

Christella 
chaseana 



r?:T 

Za.Zb.N.SA. 



Terrestrial; edges of Pinus plantation. 



X : as above 

ZP, ZS, ZT, ZD = 2 spp. 

Nephrolepis 
undulata 



Ophioglossum 
reticulatum 



a : T Terrestrial; edges of forest and plantations 

Za.Zb. M. (ZP, ZS); in grassy areas of open woodland (ZS, ZT, 

ZD). 

a : T Terrestrial; widespreading by underground 

Za.Zb.L.N.SA. stolons; at edges of forest and conifer plantations 

(ZP, ZS), woodlands (ZS), and in gardens, on Golf 
Course and lawns of the Botanic Garden, (ZT). 



X : as above 

ZP, ZS, ZT (4 spp. including 1 var. of a sp. in Habitat I). 



Dryopteris 
athamantica 



f to r : T Erect terrestrial; frequent at roadside on 

Za.Zb. L.M.S.SA. montane grassland; occasional at montane 

grassland/forest margins; rare on lower slopes. 



308 



FERN GAZETTE: VOLUME 13 PART 5 (1989) 



D. inaequalis 
var. inaequalis 
(see Habitat I 
for other var.) 


a:T 
Za.Zb.L.M.SA. 


Pteridium 
aquilinum 
subsp. aquilinum 


a to r : T 
Za.Zb.L.M.SA. 


Selaginella 
abyssinica 


a:T 
Za.Zb.M. 



Tufted terrestrial with spreading fronds; 
in moist shady margins of forest and plantations 
(ZP, ZS), woodlands (ZS), and between river 
and cultivated areas (ZS, ZT). 

Terrestrial; abundant at montane forest/ 
grassland margins, along forest paths, in 
clearings and at plantation margins (ZP, ZS); in open 
woodlands (ZS); rarely below 1000m (ZT), at edges 
of cultivation. 

Terrestrial; usually upright but sometimes 
pendant on vertical earthbanks; at woodland edges 
and plantation margins (ZP, ZS), roadside banks 
(ZS, ZT). In slight shade or open areas. 



X : as above 
ZP, ZS = 5 spp. 




Asplenium 
buettneri 


r:T 
Za.M. 


Cheilanthes 
multifida 


a:T/L 
Za.Zb.L.M.N.SA 


Hypolepis 
sparsisora 


f to a : T 
Zb.M.S.SA. 



Pellaea 
quadripinnata 

Pteris 
catoptera 



a:T 
Zb.L.M.N.S.SA. 



a to f : T 
Za.Zb.M.SA. 



Tufted terrestrial; in deep shade; thicket at 
edge of montane forest (ZP); edge of Pinus 
plantation (ZS). 

Erect terrestrial; in most exposed margins of 
Plateau, edges of forest, plantations, cliff-edges; 
lithophyte of montane grassland/forest margins 
(ZP,ZS). 

Erect but spreading terrestrial; in wet partially 
shaded marginal areas; edges of riverine forest, 
stream edges of Pinus plantations; similar habit to 
Pteridium and often mixed with it, but only in moist 
places. 

Large erect terrestrial; on rocky montane 
grassland edges, cliff-edges (ZP); conifer plantations 
margins (ZP, ZS). In open or slight shade. 

Terrestrial; spreading fronds, very variable size; 
large plants inside edges of plantations (ZP, ZS), 
small along margins. 



XI : sheltered habitats (all areas = 1 1 spp. including 1 var. of a sp. in Habitat XII). 
ZT only = 4 spp. 



Platy cerium 
elephantotis 


r (s) : E 
Za.M. 


Adiantum 
raddianum 


o:T 
Zb.M.SA. 


Aspidotis 
schimperi 


r(s):T 

Za.Zb 


Ophioglossum 
vulgatum 


f:T 

Za.Zb.SA. 


XI : as above 

ZP, ZS, ZT, ZD = 1 sp. 




Pellaea 
calomelanos 


f:T 

Zb.M. 


var. swynnertomana 





Epiphyte, in crown-base of old Jacaranda tree; 
one location. 

Terrestrial; on rocky roadside banks and in 
water channels, in slighty shady positions. 

Tufted terrestrial; at edge of brick-built 

roadside ditch; in shade of trees. First collection for 

Zomba: 5-2-85, Berrie 600 (MAL). 

Terrestrial, spreading by under-ground stolons; 
in shaded grassy area ("lawn"). 



Erect, tufted, terrestrial; edges of rocky 
outcrops, in rock crevices; on rocky roadside 
banks; usually slightly sheltered by trees or shrubs. 



PTERIDOPHYTES OF ZOMBA MOUNTAIN 



309 



XI : as above 

ZP, ZS, ZT = 2 spp. 

Adiantum 
philippense 



A. poiretii 
var. poiretii 



a to r : T 
Za.Zb.M.SA. 



o:T 
Za.Zb.B.L.S.SA. 



Pendant terrestrial; on vertical earthbanks of 
forest roads (ZP, ZS); on rocky roadsides and bridges 
(ZS, ZT); in roadside ditches (ZT); usually in slightly 
shaded places. Rare on Piateau, abundant on lower 
mountain slopes. 

Terrestrial, may be mixed with A. philippense, 

in similar habitats but usually more shaded positions. 



XI : as above 
ZP, ZS = 1 sp. 

Selaginella 
mittenii 



f : T Creeping terrestrial; in sheltered rock crevices; 

Za.Zb.L.S.SA. in moist shade below overhanging rocks. 



XI : as above 

ZS, ZT, ZD = 1 sp. (var. of this sp. in Habitat XII) 



Doryopteris 
concolor 
var. kirkii 



f:T 
Za.Zb.M.N.SA. 



(see Habitat XII for other var.) 



Erect, tufted terrestrial; in shallow soil of rocky 
places; rock crevices; on bases of trees; roadside 
banks; usually in slight shade. 



XI : as above 
ZS, ZT = 2 spp. 

Adiantum 



Pyrrosia 
schimperiana 



r:T 
Za.Zb.M.N.SA. 



a:E/L 
Za,Zb.M.SA. 



Tufted terrestrial; in rock crevices, at edge of 
woodland (ZS); between stones of roadside wall, in 
heavy shade of trees (ZT). 

Creeping epiphyte; on upper trunk and main 
branches 3-6m from ground, on all sides and 
surfaces, pendant or erect fronds; on trees of upper 
and lower mountain slopes and ZT; lithophyte of 
lower slopes and ZT. Most common epiphyte of drier 
places around Zomba Mt. 



XII : exposed habitats (all areas = 8 spp. + 1 var. = 9 spp.) 
ZP only = 1 sp. 



Mohria lepigera 



Za.Zb.M. 



Sept. 1859 Kirk (K*), "Dzomba, Zambesia 6-7000ft' 

20-5-82, Salubeni 3238 (MAL*), "on rocks in 

montane grassland" 

7-4-84, Brummitt 17130 (K, MAL*), 

"steep rocky slope" 



XII : as above 

ZT only = 1 sp. (var. of this sp. also in Habitat XI). 



Doryopteris 
concolor 
*var. nicklesii 



s 
Za.Zb. 



25-3-50, Sturgeon F2 (MAL*), 

"fully exposed rocky area" (cf. var. kirkii in Habitat 

XI.) 



310 



FERN GAZETTE: VOLUME 13 PART 5 (1989) 



XII : as above 
ZD only = 1 sp. 

*Pellaea 
longipilosa 



s" 
Za.Zb. 



14-3-77, Brummitt 14865 (K, M AL*), 
"rocky outcrop 2K E. of Zomba 



XII : as above 
ZP, ZS, ZT = 2 spp. 
Pellaea dura 

P. viridis 



f:T 
Za.Zb.SA. 



f:T 
Za.Zb.L.M.N.SA. 



Small tufted terrestrial; in b'are rocky places, 
open woodland and roadside banks; very exposed, 
extremely dry positions. 

Terrestrial, tufted but spreading fronds; on 
rocky slopes of open woodland, and roadside banks; 
sometimes mixed with P. dura but not often 
extremely exposed. 



XII : as above 




ZP, ZS, = 4 spp. 




Arthropteris 


r:T/L 


orientalis 


Za.Zb.M. 


Lycopodium 


r:T 


cernuum 


Za.Zb.M.SA. 


L. clavatum 


a to r : T 




Za.Zb.L.M.SA 



Mohria 
caffrorum 



a:T/L 
Za.Zb.L.M.SA. 



Creeping terrestrial; on rocky open slopes; 

on top of rocky outcrop, in moss/thin soil layer. 

Creeping and upright, spreading terrestrial; 

on steep earthbank at roadside; open grassy bank 

near Dam; sometimes in slight shade. 

Wide-creeping terrestrial; in wet to very dry 
places; open streamside edge of Pinus plantation, 
marshy source of river, dry edges of Eucalyptus 
plantation, dry rocky roadside bank; can occasionally 
dominate the ground flora, especially after tree- 
felling. 

Erect tufted terrestrial; on very exposed cliff-top 
grassland (ZP) and cliff-edges (ZP), and vertical 
rocky roadside (ZS). 



Local and Regional distribution 

Local distribution of species is given (a = abundant, f = frequent, o = occasional; r = 
rare; and s = single collection) for each species in each Habitat area. Some taxa have not 
been seen during the present study 1980-86. These are marked with an asterisk and local 
distribution is not given. Previous collections of these species are listed with their location 
in herbaria, using standard abbreviations (Stafleu, et al 1981). An asterisk there, e.g. K*, 
indicates that particular specimen was seen by me at that herbarium in Sept. 1986. 

Regional distribution of each species is given for the whole of Southern Africa, taken 
from the most recent available literature as follows, (abbreviations for each country in 
brackets): Zambia (Za) Kornas 1979; Zimbabwe (Zb), Burrows 1983; and all others from 
Jacobsen 1983, Botswana (B), Lesotho (L), Mozambique (M), Namibia (N), Swaziland (S) 
and South Africa (SA). . 

Plant habit 

The terms epiphyte (E), lithophyte (L), terrestrial (T), and aquatic (A), are used to describe 
plant habit, with further detailed descriptions of each species. Alternative habits showing 
the most usual habit as first letter, include E/L, E/T/L, T/E, T/L, T/A and T/L/A/ 
E. 



PTERIDOPHYTES OF ZOMBA MOUNTAIN 



311 



TABLE 2 

Plant habit. Number of species of each plant habit type in Habitats I-XII. (Abbreviations: E = epiphytic, 
T = terrestrial, L = lithophytic, A = aquatic; with alternative habits the first letter indicates the most 
usual habit. An asterisk * marks a species or variety not seen in the p-esent study, these are not 
included in the final % totals). 



Habitat, 






PLANT HABIT 








and number of 
species in each 














E 


E/L E/T/L 


T/E T//L T/L/A/E T/A 


T 


L 


A 


I: 5 spp. 
(incl. 1 sp/v 
in Habitat X) 


1 




2 


1 + 
1 sp/v 






II: 3 spp. 




2 1 










Ill: 5 spp. 


4 






1 






IV: 1 sp. 








1 






V: 12 spp. 


2 

+ r 




1* 


3 
+ 2* 


3* 




VI 8 spp. 


i 


4 1 


1 


1 






VII: 20 spp. 


i 




1 5 


12 

+ 1* 






VIII: 2 spp. 












2* 


IX: 7 spp. 






1 2 


4 






X: 14 spp. 
(13 + 1 sp/v 
in Habitat I) 






1 


12 + 
1 sp/v 






XI: 1 1 spp. 
(incl. 1 sp/v 
in Habitat XII) 


i 


1 




8 + 
1 sp/v 






XII: 9 spp. 
(8 + 1 sp/v 
in Habitat XI) 






2 

+ r 


4 


1*+ 
1 sp/v* 




total 95 spp. 83 
+ 12 spp.* 
(+ 1 sp/v*) 


= 10 

= r 


+7 +2 


+3 +4 +1 +7 
+2* 


+49 

+3* 



+4spp* 

& 
lsp/v* 



+2spp* 


% of total: 


10.5 


7.4 2.1 


3.2 4.2 1 7.4 


51.6 










E = 


20% (+1%*) 


T = 67.4% (+5.3%*) 




L = 

(+4.2%*) 


A = 
(+2.1%*) 



In some rocky habitats it is difficult to distinguish between terrestrial and lithophytic 
habit but any plant rooted in soil of more than 1cm depth is listed as T. The lithophytes 
may be in a moss layer on rock or very shallow soil layer, but they are never on a completely 
bare rock surface. 



ECOLOGY 
Montane forest pteridophytes occur in Habitats I, II, III, and VI showing that there are 



312 FERN GAZETTE: VOLUME 13 PART 5 (1989) 



a number of microhabitats or niches within montane forest in which groups of pteridophyte 
species are found together (Figure 6). 

Habitat I includes the darkest, wettest part of montane forest and only five species 
are here. There is very little ground flora other than scattered ferns. Dryopteris inaequalis var. 
atropaleacea is in very deep shade near the edge of the forest, inside the thicket of Philippia and 
Hypericum. The contrast in light intensity from the open grassland outside the forest into 
this thicket is extreme. Similarly in the deepest part of the forest, near the small stream, 
it is dark due to the dense canopy of the continuous tree-cover. The other four species 
are in this darkest part, Asplenium lobatum in one small dark area of ground, with 
A.boltonii and A.dregeanum either terrestrial or very low level epiphytes on tree bases and 
rotting tree stumps. Elaphoglossum acrostichoides is also only in one site, like A. lobatum, 
in the darkest part. It is an epiphyte on a small tree with an extremely thick canopy. 
All these species are well away from the more open pathways going through the forest. 

Habitat II and III inckide those parts where just a little bit more light penetrates the 
forest, whether in montane, riverine or cliff-edge forest. In the montane forest where the 
canopy is somewhat open because of fallen tree branches, or near pathways where tree 
saplings have been cut to keep the path open, there are many fern species which also 
occur in riverine forest since the microhabitat is similar. Much of the montane forest is 
like this, with deep moist leaf litter and permanently wet soil. The air is moist at all times 
and epiphytic ferns are most luxuriant. The filmy ferns cover tree bases and tree trunks 
up to 2 or 3cm from the ground with Asplenium mannii and A.sandersonii occurring with 
them at 1-1. 5m. The maximum number and abundance of species occurs at about 2-3m 
level where the whole surface of tree trunks is wet and dripping and covered with ferns. 

Above this level the epiphytes are more spread out on the branches, these are the species 
included in Habitat VI. The conditions on the mid to upper parts of the tree crowns of 
the montane forest are very similar to those found on trunks and crown bases in riverine 
and remnant riverine forest at lower altitudes on the mountain; they are also similar to 
some parts of the closed woodland on the mountain slopes, so the eight species in Habitat 
VI list are the most widespread of the forest/woodland pteridophyte species. In different 
locations some of these ferns occupy similar niches e.g. Pleopeltis excavata, P. macrocarpa, 
Loxogramme lanceolata and Pyrrosia rhodesiana are all epiphytes in montane and riverine 
forest, but at lower altitudes (ZS) may be lithophytes in remnant riverine forest. The position 
of the epiphytes on thre trees depends on the amount of light penetrating the canopy and 
the available water, either run-off with most at the tree base or high humidity and condensation 
in the crown-base. The filmy ferns thrive in the optimum conditions of the montane forest, 
but only survive in a few places of the riverine forest and cliff-edge forest thicket, and 
are completely absent from remnant riverine forest (ZP, ZS) and the woodlands (ZS). 

Similarly in riverine forest only (Habitat V), there are some species e.g. Selaginella 
kraussiana and Blechnum attenuatum, which thrive in the continuously moist habitat but 
require some tree-cover. They can withstand some shading but not continuous exposure. 
One small part of the riverine forest where Blotiella natalensis occurs has a very dense 
ground flora of S. kraussiana, and a nearby large tree (Rauvolfia sp.) has a complete covering 
of filmy ferns from ground level to 2m, together with Asplenium mannii, A. sandersonii and 
L. lanceolata, and looks very like the tree bases in the montane forest. But here the riverine 
forest canopy is higher and more light penetrates, although there is deep shade in some 
parts, so S. kraussiana flourishes because of the very wet soil conditions. Around the plants 
of B. natalensis the water table is almost at the surface in the dry seasons and some of 
that area is flooded during the wet season. 

Six of the species listed in Habitat V, riverine forest only, have not been seen recently 
(1980-86) and may no longer be present on Zomba Mt. The felling of Pinus patula on 
some of the sloping ground near the river has sometimes severely damaged trees closer 



PTERIDOPH YTES OF ZOMB A MOUNTAIN 3 1 3 



to the river, so these have had to be felled also. In some places along the river there has 
been very little regeneration of the riverine forest due to the slashing (cutting back of ground 
flora) to keep riverside paths clear for the tourists and trout-fishermen. This means that 
when old trees, e.g. of Halleria lucida, lose branches, fall to pieces gradually, and eventually 
rot away then the particular epiphytes on that tree may be losi, perhaps permanently from 
that area. The tree gap opens up the canopy and if no young saplings take the place of 
the old tree then what was riverine forest becomes more open remnant forest, in which 
some of the terrestrial or lithophytic ferns also cannot survive. Certainly this has happened 
with Polypodium polypodioides, an epiphytic fern, frequent to occasional in distribution 
in 1967. There are now very few trees with long, low overhanging branches stretching out 
over the water as there were then, in that part of the Mulunguzi River just below the 
Trout Ponds. The river is wider now from erosion, the undermining of the riverbank is 
partly due to slashing very close to the edge so that young ferns (Cyathea spp. and Marattia) 
are not able to get established easily by the water. 

Much of the riverside vegetation between the plantations is no longer forest or even 
remnant forest. It is just a very wet habitat, sometimes partly shaded by nearby trees in 
the Pinus plantations, so the pteridophytes along the river and streams have been listed 
in Habitat VII. These species are often standing in water during the wet season. They 
may be exposed to full sunshine, though proximity to open water usually gives a high 
air humidity. Where there was forest or remnant forest twenty years ago there is now 
in some parts just a line of tree ferns. Cyathea dregei is the most abundant and is rarely 
far from the water's edge. C. thomsonii is more scattered along the river, growing with 
C. dregei. Marattia fraxinea is the most conspicuous of the large ferns, usually rooted only 
a few metres from the water, with fronds 3-4m long spreading out almost into the conifer 
plantations alongside. Marattia is usually on the silted side of the river while the opposite 
banks have masses of Osmunda regalis, rooted between large rocks where little silting has 
occurred. All these species are in shallow water where the river is slow moving. None 
are found where the river is narrow and rapid flowing. The bed of the river changes greatly 
within short distances, from rapids to meanders, to waterfalls and narrow mini-gorges between 
large boulders, and back to wider meanders with small sandy patches on one side. 

The pteridophyte species distribution reflects these changes in different parts of the 
Mulunguzi River from the narrow source below Mulunguzi Marsh, downstream as far as 
Mulunguzi Dam. Below the Dam there is remnant riverine forest in parts, almost to the 
top of the Gorge. Here the river goes over a high waterfall through narrow cliffs. Below 
the falls there is more open rocky banks with scattered trees along the river. In the spray 
of the falls is a single location for Vittaria volkensii, a mid-level epiphyte, and close-by 
among rocks a few plants of Cheilanthes farinosa. Ten to 15m from the river, as by small 
streams on the lower slopes, are scattered small plants of Asplenium pumilum, and the 
rarer but larger Microlepia speluncae. Both are short-lived, not surviving long after the 
rains stop, because at this altitude (c. 1100m), away from the tree cover, dry conditions 
soon take over. 

Away from the river and streams, at edges of forest/woodland or forest/grassland are 
some pteridophyte species with a wide distribution on many parts of the mountain. These 
are marginal habitat species (Habitat X). The most prominent is Pteridium aquilinum 
(bracken), which can survive and spread even in poor rocky soil. In many parts of the 
Plateau it is dormant in the Pinus plantations. In years 1-3 after planting Pinus seedlings, 
bracken is cut between the young trees but after that does not grow up again due to heavy 
shading. It remains along pathways and edges of plantations, but at clear-felling 25-30 
years later it sprouts up again all over the newly felled area. Pteris catoptera also has 
a tendency to re-appear further into the felled conifer areas, but may be re-colonising rather 
than remaining dormant in the soil like the Pteridium. 



314 FERN GAZETTE: VOLUME 13 PART 5 (1989) 



In wet marginal areas Pteridium is often mixed with Hypolepis sparsisora. With similar 
growth habit they can be confused from a distance but the delicate paler green of the 
Hypolepis can be picked out in the wetter parts of this habitat on the Plateau and the 
slopes. At lower altitudes, ZT and ZD, Nephrolepis undulata is the most widespread marginal 
species in grassy/woodland areas while at higher latitudes along cliff-edges are Pellaea 
quadripinnata, Cheilanthes multifida and C. inaequalis. All these species in marginal habitats 
get some shelter from nearby trees, but can be exposed to very dry conditions at some 
time of year. Most die back during the dry season to some extent on the Plateau, some 
die back completely at lower levels, but by looking for more sheltered "favourable" marginal 
habitats it is usually possible to find some plants of these species at almost any time of 
year on some part of the mountain. 

The remaining habitats (VIII: seasonal pools, IX: seasonal flush, XI: sheltered, and 
XII: exposed) have much more extreme conditions from wet to dry seasons. There are 
no seasonal pools on Zomba Mt. but in low lying parts of Zomba District some water 
ferns may be found. Two genera (Ceratopteris and Marsilea) have been recorded, though 
I have not seen any recently, 1980-86, in the Zomba area*. Seasonal flush habitats are 
more common around the lower slopes of Zomba Mt. where rocky outcrops are found. 
These areas have smooth, rounded, granite domes with flattened tops, or small flat ledges 
on some part of the slope. Small pockets of soil on the rock quickly become soaked in 
the first rain of the season and then water-logged as seepage continues from deeper soil 
higher up the slope. The whole rock surface may then have a thin water flow over it 
for many weeks of the wet season. A short dry spell with no rain, e.g. 2 or 3 weeks in 
early December, can cause complete drying out and curling up of the soil pocket, so conditions 
can be extreme even during the wet season. 

In such places Ophioglossum spp. appear each year. O.costatum is first with the early 
flowering monocots (Platycorine, Dipcadi and Lapeirousia spp.) in the wettest parts low 
on the outcrop. Higher up the granite slope and on the flat top with deeper peaty soil 
come later species, O.gomezianum being abundant and widely distributed, and O.polyphyllum 
much rarer only in one location. In the wettest parts the later (Jan-Feb.) arrival of Isoetes 
sp. completes the O.costatum community. Another outcrop area, not so wet, has Isoetes 
with O.gomezianum and patches of Selaginella njam-njamensis. At the edges of seasonal 
flush areas clumps of Actiniopteris spp. occur. 

The seasonal flush habitat grades into the sheltered (XI) and the exposed (XII) habitats 
particularly on poorer stony soil. The terrestrial ferns Adiantum spp., Pellaea spp., Doryopteris 
concolor and Mohria spp. are all in these very dry habitats with varying amounts of shade 
or exposure. The only epiphyte which is abundant in drier places is Pyrrosia schimperiana. 

The only fern found recently in Zomba and not listed in Table 1 in Habitats I-XII 
is one which appeared amongst other ftrns in trays in the greenhouse. Some samples of 
Ophioglossum reticulatum were collected with a large amount of soil on 23-3-84 from margins 
of a Pinus patula plantation on Naisi Spur (ZS). They were planted in that soil in trays 
in the Biology Dept. greenhouse at Chancellor College, Zomba (ZD). Sometime later, in 
June, small sporelings of Pityrogramma calomelanos (L.) Link var. calomelanos appeared 
in the tray with the Ophioglossum. P. calomelanos is not previously recorded for Zomba 
and I have not found it in the wild, nor seen it in cultivation here. The specimen in MAL. 
is Berrie 422B, 7-6-84, 'weed in tray of Ophioglossum reticulatum in Biology Dept. greenhouse, 
CC It is only included here to complete the listing of all known pteridophytes from the 
Zomba area, thus the total pteridophyte taxa for Zomba, including this weed species, are 
96 species in 46 genera of 22 families, but P. calomelanos is not included in Table 1. 

*My collection closest to Zomba is M. subterranea Lepr. from Mangochi District, 100km north of 
Zomba, Berrie 316 (BM, MAL), 8-4-82; the most southern record for this species (Launert, pers. comm. 
1984). 



PTERIDOPH YTES OF ZOMB A MOUNTAIN 3 1 5 



DISCUSSION 
The 95 species listed in Table 1 include 45 genera in 22 families of pteridophytes. This 
is a relatively high number for Africa since a total of 55 species is the estimate for the 
whole of tropical Africa (Parris 1985). Thus Zomba Mt. has 19% of this total in an area 
of approx. 8 x 12km. This compares with the Zambia records (Kornas 1979) where the 
highest number of pteridophyte species in a one degree square (l°lat x l°long., approximately 
100km x 100km) is 56 species or 11% of the total for Africa. The Lusaka District at 
approximately the same latitude as Zomba, but 800km due^west at longitude 28-29°E, has 
31 species, only 6.2%. Lusaka, the Zambian capital, at 1278m altitude has only 835mm 
mean annual rainfall compared to Zomba (alt. 900m) with 1464mm, and Zomba Plateau 
(alt. 1900m) with 2244mm mean annual rainfall. This big difference in rainfall is probably 
the main cause of the big difference in number of pteridophyte species between Lusaka 
and Zomba. 

Similarly on Zomba Mt. the differences between the Plateau rainfall and the Town 
rainfall is reflected in the .plant distribution and particularly in the distribution of the 
pteridophyte species. The montane forest is only on Zomba Plateau and most of the riverine 
forest is also on the Plateau. The pteridophyte numbers in these forests are 21 spp. in 
montane and 29 spp. in riverine, with 16 spp. common to both (Figure 6b). Of the 68 
spp. in all habitats (Fig. 6e) on Zomba Plateau there are 32 forest/woodland species with 
22 of these restricted to ZP. (Fig. 6a & b). The number of forest/woodland species, like 
the rainfall, decreases with altitude from ZP, 32; ZS, 10; ZT, 5 and ZD, 2 spp. The marginal 
habitat species follow the same pattern with ZP, 13; ZS, 12; ZT, 6 and ZD, 2 spp. The 
riverside non-forest (VII) habitat has a slightly different pattern with ZP, 11; ZS, 16; ZT, 
10 and ZD, 4 spp. and has discontinuous distribution (3 spp.) also, not found in the other 
habitats. This reflects the particular microhabitat required by these three species rather 
than being related to altitude or rainfall. This is shown by Osmunda regalis on ZP, ZS 
and in ZD. The Mulunguzi River in ZT has rapid flowing water, mostly a narrow riverbed 
with huge rocks and boulders at each side. Osmunda grows on sandy, silted banks or 
between small rocks in slower water so it is not found in ZT. Osmunda is also only by 
the main river, not by small streams which dry out completely in the dry season. Cyclosorus 
interruptus is similar with swampy riverine area (ZP), muddy streamside (ZT) and silted, 
swampy stream (ZD) being the microhabitats in three areas, but no suitable places occur 
on ZS for it. Amauropelta bergiana (ZP and ZT) may be undercollected, since the genera 
of Thelypteridaceae, being very alike, need further investigation ecologically. Christella 
chaseana (Habitat X: marginal, ZS) in the same family is also queried (?r) in distribution, 
needing further collection. 

The flush habitat (IX) and sheltered habitat (XI) have similiar patterns with more species 
on the mid-mountain slopes and township area (ZS, ZT), while the exposed habitat (XIII) 
reverts to the pattern of Habitats I-VI. Taking these all together (Fig. 6e) gives a clear 
picture of greater numbers of species at higher altitudes, ZP 68 spp. (72% of total spp.), 
ZS 55 spp. (58%), ZT 39 spp. (41%) and ZD 15 spp. (16%). The factors affecting this 
distribution due to altitude are the increased rainfall at higher altitudes; greater tree-cover 
i.e. thicker forest on the higher land; riverine and remnant riverine on ZP and ZS; woodland 
on upper and lower slopes; and the amount of condensation-cloud on the Plateau and 
upper slopes especially during the dry season. The higher temperature at lower altitudes, 
with the very strong dry winds of mid-dry season (Aug.), will also combine with the lack 
of tree-cover, in ZD, to give a smaller number of species. Only in streamside habitat can 
they survive the dry season, if the stream is a permanent one. 

An analysis of the plant habit of species in the different habitats (Table 2) shows 20% 
of the 95 species are usually epiphytic though 9.5% may occasionally be lithophytic or 
terrestrial. 51.6% of the total are always terrestrial while 7 spp. (7.4%) are so close to 



316 FERN GAZETTE: VOLUME 13 PART 5 (1989) 



water as to be sometimes actually in it, classed as T/A. Only 2 spp. (2.1%) are true aquatic 
ferns. 

Habitat V (riverine forest only) has suffered the most since the 1960's if the species 
marked are not any longer present on Zomba Mt. These species are 3 lithophytic, 2 terrestrial, 
1 T/L, and 1 epiphyte all previously recorded from riverine areas. During the 1980-86 
period I have made observations on the pteridophyte flora of most parts of Zomba Mt. 
I know all these species from other parts of Malawi and it is possible they may be on 
the mountain, in some place where I have not seen them, if so I would regard them as 
threatened species. 

ACKNOWLEDGEMENTS 
I would like to thank Dr J.H. Seyani, Curator of the National Herbarium of Malawi, 
for advice and discussion on my work during the preparation of this paper, also the staff 
of the Herbarium, especially Mr I.H. Patel and Mr E. Tawakali, for their assistance in 
fieldwork on Zomba Mt. 

Thanks are also due to the Research and Publications Committee of the University 
of Malawi for support from research Grant No. 8212, 1982-86 for transport for fieldwork. 

Finally I would like to thank Dr B.S. Parris and Mr P.J. Edwards for help, discussion 
and encouragement during my very brief visits to the Kew Herbarium in Aug. 1984 and 
Sept. 1986. 

REFERENCES 
BAKER, C.A. 1959. A map of Shire Valley and Highlands Routes of Livingstone, 1859. Nyas Journ. 

12(2): 22-26. 
BALLARD, F. 1953. Pteridophyta. In J.P.M. Brenan et al. Plants collected by the Vernay Nyasaland 

Expedition of 1946. Mem. N. York Bot. Gdn. 8(3): 197-210. 
BERRIE, A. 1984. A checklist of the pteridophytes of Zomba Mt., Malawi. Luso: J.Sci. Tech. (Malawi) 

5(2): 67-86 
BRASS, L.J. 1953. Vegetation of Nyasaland. Report on the Vernay Nyasaland Expedition of 1946. 

Mem. N. York. Bot. Gdn. 8(3): 161-190. 
BURKILL, I.H. 1906. List of the known plants occurring in British Central Africa, Nyasaland, and 

the British Territory north of the Zambesi. Botanical Appendices. Appendix II. pp. 233-284 

and 284a-2841 in H.H. Johnston British Central Africa. (3rd Ed.) Methuen, London. NOTE: 

pp. 284a-2841 the Supplementary List by Burkill, are only in the 3rd Edition 1906, not in the 

1st Ed. 1897, nor in the 2nd Ed. 1898, by Methuen. The 1969 Reprint of the 1st Ed. by Negro 

Universities Press, New York also does not have the Supplementary List, pp. 284a-2841. 
BURROWS, J.E. & S.M. 1983. A checklist of the pteridophytes of Zimbabwe. J. S. Afr. Bot. 49(3): 

193-212. 
C.C. : Geography Dept., Chancellor College, Zomba. Meterological Records 1976-86, (unpub.) 
F.R.I.M. : Forestry Research Institute of Malawi, Zomba. Meterological records for Zomba Plateau. 

Rainfall data 1976-86; Temperature data 1978-80 (unpub.) 
HOLTTUM, R.E. 1974. The Thelypteridaceae of Africa and adjacent islands. J. S. Afr. Bot. 40(27: 

1923-168. 
JACKSON, G. 1959. Correspondence of John Kirk to Sir William Hooker, Dr J. Hooker, and Professor 

Balfour. Nyas. Journ. 12(2): 40-54. 
JACOBSEN, W.B.G. 1983 The Ferns and fern Allies of Southern Africa. Butterworths, Durban, S. 

Africa. 542 pp. 
KORNAS, J. 1979. Distribution and ecology of the Pteridophytes in Zambia. Warszawa-Krakow, 

Panstwowe Wydawnictwo Naukowe. 206 pp. 
PARRIS, B.S. 1985. Ecological aspects of the distribution and speciation in Old World tropical ferns. 

Proc. Roy. Soc. Edinb. 86B: 341-346. 
SCHELPE, E.A.C.L.E. 1970. Pteridophyta In A.W. Exell & E. Launert (Eds) Flora Zambesiaca. Crown 

Agents, London. 254 pp. 
STAFLEU, F.A. HOLMGREN, K.H., KEUKEN, W., & SCHOFIELD, K. 1981. Index Herbariorum 

Pt. 1, 7th Ed. Bohn, Scheltema & Holkema, Utrecht. 452 pp. 
WALTER, H. 1971. Ecology of Tropical and Subtropical Vegetation. Oliver & Boyd, Edinburgh. 

539 pp. 
WHITE, F. 1983. The vegetation of Africa. A descriptive memoir to accompany the UNESCO/AETFAT/ 

UNSO vegetation map of Africa. No. XX in series 'Natural Resources Research' UNESCO, 

Paris. 356pp. 



FERN GAZ. 13(5) 1989 317 



PRELIMINARY REPORT OF CHROMOSOME COUNTS IN THE 
GENUS AZOLLA (PTERIDOPHYTA) 

KALLIROI K. STERGIANOU and KEITH FOWLER 

School of Biological Sciences, Portsmouth Polytechnic, King Henry I Street, 
Portsmouth POl 2DY, Hampshire, England 

ABSTRACT 

Contrary to previous reports, study of chromosome numbers in the genus Azolla establishes 
the occurrence of 2n=44 in all species except A. nilotica which has 2n=52. In addition, 
triploid clones (2n=66) were found in some species and a single tetraploid clone of A. 
pinnata (2n=88) is recorded for the first time in the genus. 

INTRODUCTION 

Azolla Lam. is a genus of heterosporous aquatic ferns with a world-wide distribution in 
tropical to temperate regions. Five species are currently recognised within section Azolla 
: A. filiculoides Lam., A. rubra R.Br., A. caroliniana Willd., A. mexicana Presl and A. 
microphylla Kaulf.; and two species in section Rhizosperma (Mey.) Mett. : A. nilotica Decne. 
ex Mett. and A. pinnata R. Br. Chromosome numbers are recorded for most species but 
show contradictory results, and are often based on a single clone (Litardiere 1921, Tschermak- 
Woess & Dolezal-Janisch 1959, Loyal 1958, Loyal, Gollen & Ratra 1982, Thanh 1983, 
Singh, Patra & Nayak 1984, Lin & Sleep 1988). 

MATERIALS AND METHODS 
Extensive investigation of somatic chromosome numbers in Azolla is in progress at 
Portsmouth Polytechnic. Approximately eighty clones of all species from widely separate 
regions within the geographical range of the genus have been studied (Table 1). It should 
be mentioned, however, that only two clones of A. nilotica were obtained, and more material 
will need to be examined. The large number and small size of chromosomes in Azolla 
indicate that meiotic rather than somatic material would be easier to examine. Vegetative 
material was used, however, because it is generally considered more appropriate in establishing 
reliable chiomosome numbers in unknown taxa. Shoot tips were pretreated in 0. 1% colchicine 
for 5 hours and transferred to 3:1 absolute alcohol: glacial acetic acid for at least 24 hours. 
The material was then hydrolysed in 1 M HC1 at 60°C for 7 mins., stained with Feulgen 
for 1 hour and squashed in acetocarmine. Voucher herbarium specimens will be deposited 
at the British Museum (Natural History). 

TABLE 1 

Number of clones of species studied and their distribution range. 

Geographical distribution (* species introduced) 

Europe *; North, Central & South America; East Asia *. 

Australasia. 

Europe *; Central Africa *; North, Central & South America. 

North & South America; East Asia *. 

Central & South America. 

East Africa. 

West, East, Central & South Africa; India; Southeast Asia; East Asia; 
Australasia. 



Species 


Clones 


A. filiculoides 


13 


A. rubra 


2 


A. caroliniana 


16 


A. mexicana 


9 


A. microphylla 


13 


A. nilotica 


2 


A. pinnata 


26 



318 



FERN GAZETTE: VOLUME 13 PART 5 (1989) 






* 



tH .HI 



• • *♦ 



&$ % % 



4m sum 



4 



* * 

FIGURE 1. Mitotic metaphases (Scale bar = 3 jam). 

a) A. filiculoides showing 2n=44. (Portsmouth Polytechnic Coll. No. 103, UK) 

b) A. caroliniana showing 2n=44. (IRRI Coll. No. 3002, USA) 

c) A microphylla showing 2n=44. (IRRI Coll. No. 4018, Paraguay) 

d) Triploid A. pinnata showing 2n=66. (IRRI Coll. No. 5, Thailand) 



AZOLLA CHROMOSOME COUNTS 319 



RESULTS AND DISCUSSION 
Despite previous reports to the contrary, the results indicate that the chromosome number 
is invariable and will provide little information useful for taxonomic separation, particularly 
in section Azolla. With the exception of A. nilotica, which was found to be 2n=52, counts 
of 2n=44 were recorded from clones of all the other species (Fig. la, b and c). Most species 
in section Azolla appear to have similar chromosome size. Although A. pinnata (section 
Rhizosperma) has the same chromosome number as species of section Azolla, it has distinctly 
smaller chromosomes. Triploid counts (2n=66) were recorded in both sections of the genus. 
In section Azolla, triploidy was found in only one clone of A. filiculoides (from Bolivia) 
and A. microphylla (from Mexico), and in two clones of A. caroliniana (from Brazil and 
Mexico). In section Rhizosperma triploids appear to be more frequent, being found in five 
clones of A. pinnata from Nigeria, Sri Lanka, Thailand, China and Australia (Fig. Id). 
The incidence of tetraploidy (2n=88) was low, being found in only a single clone of A. 
pinnata from Bangkok. Further investigations aim to clarify whether the triploid clones 
have resulted from hybridisation between tetraploid and diploid clones, or are produced 
by the occasional function of unreduced gametes. Functional unreduced gametes are usually 
derived from interspecific hybrids, but can also be produced from species clones under 
certain environmental conditions. Further meiotic or electrophoretic studies of these triploids 
may establish whether they are auto- or allotriploids, and could provide evidence for their 
origin. It remains to be discovered whether the clone of A. pinnata represents a tetraploid 
population or is simply a unique clone, and if tetraploid populations occur. 

ACKNOWLEDGEMENTS 
We are grateful to the Overseas Development Administration for supporting the Azolla 
research programme at Portsmouth (Grant R 4359 awarded to K. Fowler, Sept. 1987). 
Our thanks also to Dr I. Watanabe of IRRI (International Rice Research Institute, Philippines) 
for providing most of the material for study, and to Colin Derrick (School of Biology) 
for photographic work. 

REFERENCES 
LIN, Y.X. & SLEEP, A. 1988. A chromosome count from Azolla filiculoides (Azollaceae: Pteridopbyta). 

Fern Gaz. 13(4): 193-198. 
LITARDIERE, R. de. 1921. Recherches sur l'element chromosomique dans la caryocinese somatique 

des Filicinees. Cellule 31: 255-274. 
LOYAL, D.S. 1958. Cytology of two species of Salviniaceae. Curr. Sci. 27(9): 357-358. 
LOYAL, D.S., GOLLEN, A. K. & RATRA, R. 1982. Morphological and cytotaxonomic observations 

on Azolla pinnata. Fern Gaz. 12(4): 230-232. 
SINGH, P.K., PATRA, R.N & NAYAK, S.K. 1984. Sporocarp germination, cytology and mineral 

nutrition of Azolla species. In SILVER, W.S. & SCHRODER, E.C. Practical applications of 

Azolla for rice production. Proc. Intern. Workshop, Mayaguez, Puerto Rico, Nov. 17-19, 1982: 

55-65. Nijhoff-Junk. 
THANH, LE DUY. 1983. Cytology and morphogenesis of sexual organs of Azolla. Intern. Congress 

of Genetics, New Delhi, Dec. 12-21, 1983: 699. 
TSCHERMAK-WOESS, E. and DOLEZAL-JANISCH, R. 1959. Uber die karyologische Anatomie 

einiger Pteridophyten sowie auffallende Unterschiede im Kernvolumen bei Cyrtomium falcatum. 

Ost.Bot.Z. 106:315-324. 



320 FERN GAZETTE: VOLUME 13 PART 5 (1989) 



THE FERN GAZETTE 



NOTES FOR CONTRIBUTORS 

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The British Pteridological Society 
THE FERN GAZETTE 

VOLUME 13 PART 5 1989 

CONTENTS 



Page 

MAIN ARTICLES 

The history of Diphasiastrum issleri (Lycopodiaceae) in Britain 

and a review of its taxonomic status 

- AC Jermy 257 

Compression and slingshot megaspore ejection in Selaginella selaginoides 
A new phenomenon in pteridophytes 

- CM Page 267 

An aberrant form of Equisetum telmateia (Pteridophyta) 
from the West of Ireland 

- Margaret R.I. Westwood 277 

A new species of Selaginella (Pteridophyta) from Cameroon, West Africa 

- Nat Quansah 282 

New ferns of Madeira 

- Mary Gibby & J.D. Lovis 285 

The ecology and distribution of pteridophytes of Zomba Mt., Malawi 

- Audrey Berne 291 
Preliminary report of chromosome counts in the genus Azolla (Pteridophyta) 

- Kalliroi K. Stergianou & Keith Fowler 317 



SHORT NOTE 

Dryopteris x fraser-jenkinsii - a correction 

- Mary Gibby & C.-J. Widen 276 

REVIEWS 266 

281 



(THE FERN GAZETTE Volume 13 Part 3 was published on 28th July 1987 
Part 4 on 15th October f9,88) 



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ISSN 0308—0838 

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SOCIETY 



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Volume 13 Part 6 



1990 



THE BRITISH PTERIDOLOGICAL SOCIETY 
Officers and Committee for 1990 

President: Dr Barry A. Thomas 
President Emeritus: James W. Dyce 

Vice-Presidents: James A. Crabbe, Dr R.E. Holttum, A. Clive Jermy, 

Reginald Kaye, Gwladys Tonge 

Honorary General Secretary and A.R. (Matt) Busby 

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(Tel: Coventry 715690) 

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Department of Botany, The Natural History Museum, 
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Treasurer: Dr Nicholas J. Hards, 

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Meetings Secretary: Anthony C. Pigott, 

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The Natural History Museum, Cromwell Road, London SW7 5BD 

Editor of the Pteridologist: Martin H. Rickard, 

The Old Rectory, Leinthall Starkes, Ludlow, Shropshire, SY8 2HP 

assisted by J.W. Dyce 

Committee: Patrick J. Acock, Peter G. Barnes, Jack H. Bouckley, Josephine M. Camus, 

Christopher R. Fraser- Jenkins, Jennifer M. Ide, Margaret Nimmo-Smith, 

R. Neil Timm, Dr Trevor G. Walker, John R. Woodhams 

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102 Queens Close, Harston, Cambs., CB2 5QN 

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201 Chesterton Road, Cambridge, CB4 1AH 

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The BRITISH PTERIDOLOGICAL SOCIETY was founded in 1891 and today continues as a focus 
for fern enthusiasts. It provides a wide range of information about ferns through the medium of its publications 
and available literature. It also organises formal talks, informal discussions, field meetings, garden visits, 
plant exchanges, a spore exchange scheme and fern book sales. The Society has a wide membership which 
includes gardeners, nurserymen and botanists, both amateur and professional. The Society's journals, the 
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FERN GAZ. 13(6) 1990 321 



SCANNING ELECTRON MICROSCOPE PHOTOGRAPHS OF THE 

SORI AND SPORES OF SIX SPECIES OF RUST FUNGI (UREDINALES) 

FOUND ON FERNS IN BRITAIN 

ADRIAN J. HICK & THOMAS F. PREECE 
Department of Pure and Applied Biology, The University, Leeds LS2 9JT, U.K. 

ABSTRACT 
Scanning electron microscope photographs are presented of Dryopteris dilatata and Abies 
alba with the rust Milesina kriegeriana; Blechnum spicant and Abies alba with Milesina 
blechni; Asplenium scolopendrium and Abies alba with Milesina scolopendrii; Polypodium 
interjectum with Milesina dieteliana; Phegopteris connectilis with Uredinopsis filicina; 
and Cystopteris fragilis with Hyalopsora polypodii. In each case a sorus of the rust 
and individual spores at high magnification are shown. A table is included of other 
British ferns on which rust fungi have been recorded. The need for more information 
about the occurrence and distribution of these rusts on ferns in Britain is emphasised. 

INTRODUCTION 
A review of the literature in relation to an introductory S.E.M. atlas of sori and spores 
of some rusts (Preece and Hick 1989) and many discussions with other biologists in relation 
to rust fungi clearly indicated two things in relation to rusts on ferns. No British plant 
pathologist we discussed the matter with was aware that there were rusts on ferns, and 
eminent pteridologists were equally uncertain as to whether they occurred, or not! A thorough 
review of the literature and discussion of the fern rusts is long overdue. 

This is not such a review, but a presentation of S.E.M. photographs of six fern rusts 
collected in Britain, and of which no S.E.M. photographs exist in the literature. 

Rust workers have thought rusts on ferns to be the primitive forms from which other 
rusts, (on angiosperms), have evolved (Ando 1984). An important recent cladistic analysis 
suggests however that this is not the case, and that tropical short-cycled rusts (usually with 
only urediniospores) are the basal group in the evolution of rust fungi (Hart 1988). We 
do not ascribe any specific biological usefulness, hypothesize about phylogenetic connections, 
or speculate about the beautiful surface ornamentation visible in the S.E.M. pictures presented 
in this brief paper. We hope that these pictures will encourage others to consider these 
matters. It might be that (as with other rust fungi on other plants) the extreme host specificity 
of the rusts is of taxonomic value to pteridologists? At the very least we hope to encourage 
British fern-workers to look out for rusts and thus enable our knowledge of the occurrence 
and distribution of rust fungi on ferns in the British Isles to expand. We would welcome 
specimens from any location, pressed and dry (not in plastic bags). 

According to Peterson (1974) the general rust fungus life cycle is the "most plastic and 
complex" series of events known for any kind of living organism. Only the life cycles of 
certain animal parasites approach the complexity of those of rust fungi on plants, and 
quite an effort is necessary to grasp both the sequence of events in the life cycle of any 
rust fungus, the function of the various stages in the life cycle, and the different spores 
of each species which occur. An explanatory outline of these matters of rust form and 
function is given in Preece & Hick (1989), together with references enabling the reader 
to find light microscope drawings of most of the rusts on plants (including ferns) found 
in Britain. 

All the five types of rust spores: teliospores borne on telia; basidiospores produced 
from the basidia on teliospores; spermatia borne on spermagonia; aeciospores borne in 
aecia and urediniospores borne on uredinia, may be produced on the same plant host, 
or on two plant hosts. In the latter case, the urediniospores and teliospores are usually 
found on one host and the spermatia, with the aeciospores, on the other host. Some spore 
types may be missing from particular rusts. These missing stages either do not exist, or 



322 



FERN GAZETTE: VOLUME 13 PART 6 (1990) 



have not yet been found. Where an alternate host is known for any British fern rust, it 
is a gymnosperm - a conifer, of the genus Abies. The teliospores of fern rusts, where 
they are known, are enclosed within the epidermis and in some cases the leaf mesophyll 
cells and cannot be illustrated by scanning electron microscopy. More microscopic studies 
of the production of basidiospores from the teliospores of fern rusts are needed, and also 
of the spermatia which precede the production of aecia on fir needles. 

There are very few scanning electron microscope pictures of fern rust sori or spores 
in the literature. Littlefield & Heath (1979) included S.E.M. pictures of the urediniospores 
of Uredinopsis osmundae (which occurs in the USA on Osmunda spp.) showing their smooth 
surfaces with an erect wall-lTke irregular thickened ridge running along opposite sides of 
the spore, and also S.E.M. pictures of urediniospores of Milesia laeviuscula (which according 
to Arthur 1934 occurs in the U.S.A. on Polypodium californicum and P. glycyrrhiza) which 
are completely smooth. Littlefield & Heath used their pictures to illustrate the range of 
surface ornamentation (or lack of it) on the urediniospores of rusts in general. 

Using S.E.M., Hafellner & Grill (1982) have shown that the urediniospores of Milesia 
vogesiaca on Polystichum aculeatum and P. lonchitis are not smooth, as was thought to 
be the case by light microscopy, but finely verrucose. These authors confirmed, however, 
that the urediniospores of the rust Milesia whitei on P. aculeatum were typically echinulate. 



List of British ferns, by Family, and the 

OPHIOGLOSSACEAE 

OSMUNDACEAE 

ADIANTACEAE 

Adiantum capillus-veneris: 

HYMENOPHYLLACEAE 

POLYPODIACEAE 

Polypodium vulgare (sensu lato): 

HYPOLEPIDACEAE 

THELYPTERIDACEAE 

Phegopteris connectilis: 

ASPLENIACEAE 

Asplenium scolopendrium: 
A. adiantum-nigrum: 
A. ruta-muraria: 

ATHYRIACEAE 

Gymnocarpium dryopteris: 
Cystopteris fragilis: 

ASPIDIACEAE 



TABLE I 

species of rust fungi so far recorded on them* 
No rusts recorded in Britain 
No rusts recorded in Britain 

Hyalopsora adianti-capilli-veneris 
No rusts recorded in Britain 

Milesina dieteliana 

No rusts recorded in Britain 

Uredinopsis filicina 

Milesina scolopendrii 
Milesina magnusiana 
Milesina murariae 

Hyalopsora aspidiotus 
Hyalopsora polypodii 



Polystichum aculeatum: 
Polystichum setiferum: 

Dryopteris filix-mas: 


Milesina vogesiaca 
Milesina whitei 
Milesina vogesiaca 
Milesina carpatorum 
Milesina kriegeriana 


Dryopteris affmis (sensu lato): 


Milesina kriegeriana 


Dryopteris carthusiana: 


Milesina kriegeriana 


CHNACEAh 

Blechnum spicant: 


Milesina blechni 






* Fern family and species nomenclature from Clapham, Tutin & Moore (1988). Fern rusts according 
to Wilson & Henderson (1966) and Ellis & Ellis (1985) 



RUST FUNGI ON BRITISH FERNS 



323 




a 



mifim 











FIGURE 1: Dryopteris dilatata with the rust Milesina kriegeriana: a) fern sporangia and rust sori 
to show relative sizes on frond on brown necrotic area; b) rust sori at higher magnification; c) rust 
urediniospore seen below stoma; d) single rust urediniospore. Abies alba with the rust Milesina kriegeriana: 
e) aecia on lower surface of needle; aeciospores from aecium. 



324 



FERN GAZETTE: VOLUME 13 PART 6 (1990) 



Using carbon replicas, Henderson & Prentice (1977) showed the echinulations on the 
urediniospores of Milesina blechni on Blechnum spicant to develop in the same way as 
those of the urediniospores of other rust fungi. Spines arise within the primary urediniospore 
wall, followed by the deposition within it of a thicker secondary wall. This secondary spore 
wall becomes the spore wall normally seen, with the spines exposed by loss of the primary 
wall. A list of the ferns occurring in the British Isles according to Clapham, Tutin & Moore 
(1988) is given in Table 1, together with the names of the rusts which occur on fifteen 
of these fern species. Out of eleven families of ferns, four: the Ophioglossaceae, Osmundaceae, 
Hymenophyllaceae and Hypolepidaceae have no rusts recorded on them in Britain. Four 
ferns, Dryopteris filix-mas, D. affinis (sensu lato), D. carthusiana and D. dilatata, have 
the same rust (Milesina kriegeriana) on them. Two ferns, Polystichum aculeatum and 
Polystichum setiferum, have the same rust recorded on them (Milesina vogesiaca). Two 
ferns have two rusts on each species (Polystichum setiferum with Milesina whitei and M. 
vogesiaca; Dryopteris filix-mas with Milesina carpatorum and M. kriegeriana). Almost all 
the Figures in this paper are of Yorkshire fern rust specimens, and the material was prepared 
from herbarium specimens (some very old) as described in Preece & Hick (1989). 






FIGURE 2: Blechnum spicant with the rust Milesina blechni: a) portion of fern frond with two 
rust sori on brown area; b) two urediniospores from sorus. Abies alba with the rust Milesina blechni: 
c) lower surface of needle with aecia; d) aeciospores from aecium. 



RUST FUNGI ON BRITISH FERNS 



325 







f 

FIGURE 3: Asplenium scolopendrium with rust MUesina scolopendrii: a) portion of parallel-sided 
brown area of frond with two rust sori; b) two urediniospores from one of these son. Abies alba 
with rust MUesina scolopendrii: c) aecium on lower surface of needle; d) aeciospores from this aecium. 
Polypodium interjection with rust MUesina dieteliana: e) three rust sori on yellow & brown patch 
on frond; f) two urediniospores from one of these sori. 



326 



FERN GAZETTE: VOLUME 13 PART 6 (1990) 




..r-l«: 



' 



a 



■m 








FIGURE 4: Phcgopteris connectilis with rust Uredinopsis filicina: a) brown portion of frond with 
two rust sori; b) single thick-walled urediniospore ("amphispore"); c) rust sorus showing elongated 
urediniospores with apical points; d) one of these normal elongated urediniospores alongside an 
amphispore, for comparison. Cystoptcris fragilis with rust Hyalopsora polypodii: e) two uredinia from 
brown portion of frond; two urediniospores from one of these sori. 



RUST FUNGI ON BRITISH FERNS 327 



COMMENTS & DISCUSSION ON THE S.E.M. FIGURES 
FIGURE 1. Dryopteris dilatata - Abies rust: Milesina kriegeriana 

All spores are colourless, and often appear like white dust near sori on brown area on 
fronds. Uredinia shown in the Figure also occur on Dryopteris filix-mas, D. carthusiana 
and D. affmis (sensu lato). The echinulate urediniospores emerge via the centrally placed 
stoma. 

On several Abies species, including A. alba, the inconspicuous spermagonia occur most 
commonly on the upper surfaces of the fir needles, and the aecia occur as shown here 
in two irregular rows on yellowish parts on the lower surface of the needle. The aeciospores 
on the fir have distinctly different verruculose ornamentation from the echinulate 
urediniospores on the ferns. The obscure teliospores occur within the epidermal cells. There 
are relatively few (easily accessible) records of the occurrence of this common rust e.g. 
Clark (1980) on D. filix-mas from Whichford Wood close to the Warwickshire/Oxfordshire 
border; Bramley 1985 records it from High Force, Teesdale on D. dilatata. See also Preece 
& Hick, 1989, Atlas species 27. (Sometimes the name Milesia is used for Milesina, as in 
this paper. Wilson & Henderson 1966 point out that Milesia is the correct term when 
telia have not been described, and Milesina when the teliospores have been described). 
FIGURE 2. Blechnum spicant - Abies rust: Milesina blechni 

Very similar to M. kriegeriana but host-specific to B. spicant. Aecial stage recorded on 
A. alba and A. cephalonica. No recent Warwickshire records (Clark 1980). Bramley (1985) 
gives only one recent Yorkshire record - near Pickering, 1930. There are older records 
on A. pectinata and A. amabilis. As in all British fern rusts, it almost certainly occurs 
more widely. Wilson & Henderson (1966) say it is "frequent but overlooked" in Britain. 
See also Preece & Hick, 1989, Atlas species 26. 

FIGURE 3. Asplenium scolopendrium - Abies rust: Milesina scolopendrii 
Similar to other Milesina spp. illustrated here. The stage on the alternate host Abies alba 
seems very rare. There is a record on the fern from Forden in Montgomeryshire in Grove 
(1913) and it is plentiful on Hart's-tongue Fern on Llanymynech Hill, Shropshire, in 1989 
(T.F.P.). 

Though there are no records in Bramley (1985), it has recently been found in Yorkshire 
(Preece & Hick 1989). One record has been given by Clark (1980) for Warwickshire. Wilson 
& Henderson (1966) regarded this as a "scarce" rust in Britain. Characteristically, parallel- 
sided brown-black areas are seen on leaves. In these the raised uredia can be seen as illustrated. 
See also Preece & Hick 1989, Atlas species 27A. 
Polypodium inter jectum - Abies rust: Milesina dieteliana. 

As fern taxonomy develops, the rusts of ferns need closer study in the field. Anglesey and 
Cornish specimens of Polypodium vulgare (sensu lato) examined in 1989 by one of us 
(T.F.P.) are certainly Polypodium interjectum in the sense of Camus & Jermy (1987). Again 
regarded by Wilson & Henderson (1966) as a "frequent but overlooked" fern rust species 
in Britain. The Abies stage is either non-existent or very rare. Artificial inoculation experiments 
have produced aecia on A. alba and A. concolor (Wilson & Henderson 1966). 

Survey work on the different segregates of the Polypodium vulgare species could be 
most rewarding. 

FIGURE 4. Phegopteris connectilis rust: Uredinopsis filicina 

The rust genus Uredinopsis has long been considered to be the most primitive genus of 
the Uredinales, though this is now disputed (vide supra). In general in the genus, the spore 
walls and spore contents are colourless, and (as distinct from Milesina) the obscure teliospores 
occur in the leaf mesophyll. The uredinia open by tearing of the epidermis (see Figure) 
as distinct from the stomatal opening as in Milesina. There are often two types of 
urediniospore, thick walled "amphispores" often polygonal and rounded, and elongated 
urediniospores with a mucronate tip (see Figure). Uredinopsis filicina has not been recorded 



328 FERN GAZETTE: VOLUME 13 PART 6 (1990) 



on Abies in Britain, although it occurs elsewhere on this tree. There have been no Yorkshire 
records since the 1930's (Bramley, 1985). This is possibly a very rare rust (Wilson & Henderson 
1966). See also Preece & Hick (1989) Atlas species 129. 
Cystopteris fragilis rust: Hyalopsora polypodii 

These pictures are of the third and last rust genus so far found in Britain on ferns - Hyalopsora. 
This last genus is generally similar to Milesina (see Figure), except there is a pigment in 
the spore cytoplasm so the spores are yellow when fresh. The obscure teliospores are found 
in the epidermal cells only and germinate there without winter rest to produce baskfrospores. 
In Yorkshire, at the end of the 19th century, this rust was very common on the fern host 
in the "fern-cases" beloved of Victorians, as well as in the wild. 

It is now it seems rare in the Yorkshire Pennines (Bramley 1985). Grove (1913) drew 
his illustrations of the fungus from a collection made at Shrewsbury in Shropshire. It seems 
to be very uncommon in Britain on the Brittle Bladder Fern - but it needs looking for 
in glasshouses and gardens as well as in the wild. Like Uredinopsis, Hyalopsora has thick 
walled "amphispores" as well as the thin walled normal urediniospores shown on the Figure. 
It seems that the teliospores have not been seen in ferns in Britain, nor has any alternate 
aecial stage been found on Abies. Wilson & Henderson (1966) point out that in general 
world wide terms H. polypodii has a very wide host distribution and is known on "at 
least 25 species of fern in 13 genera". See also Preece & Hick (1989), Atlas species 11. 

REFERENCES 
ANDO, K. 1984. Phylogeny of the fern rusts (Uredinopsis, Milesina and Hyalopsora. 

Trans. my col. Soc. Japan 25: 295-304. 
ARTHUR, J.C. 1934. Manual of the Rusts of the United States and Canada. Purdue Research Foundation, 

Lafayette, Indiana, U.S.A., page 7. 
BRAMLEY, J.C. 1985. A fungus Flora of Yorkshire. Leeds: Yorkshire Naturalists Union, page 192. 
CAMUS, J & JERMY, C. (1987) The BM Fern Crib. London: British Museum (Natural History). 
CLAPHAM, A.R., TUTIN, T.G. & MOORE, D.M. (1988) Flora of the British Isles, 3rd Edn. Cambridge: 

University Press. 
CLARK, M.C. 1980. A fungus Flora of Warwickshire. Birmingham Natural History Society; published 

by the British Mycological Society, London, page 205. 
ELLIS, M.B. & ELLIS, J.P. 1985. Microfungi on Land Plants. London: Croom Helm. 
GROVE, W.B. (1913) British Rust Fungi. Cambridge: University Press, page 378. 
HAFFELLNER, J. & GRILL, D. 1982. Scanning electron microscopy investigations: Milesia vogesiaca 

and M. whiteii. Plant Systematics and Evolution 141 (1): 23-30. 
HART, J. A. 1988. Rust fungi and host plant co-evolution: Do primitive hosts harbor primitive parasites? 

Cladistics 4: 339-366. 
HENDERSON, D.M., & PRENTICE H.T. 1967. The morphology of fungal spores: Milesina blechni. 

Notes from the Royal Botanical Garden, Edinburgh 35: 115-117. 
LITTLEFIELD, M.J. & HEATH, M.C. 1979. Ultrastructure of rust fungi. New York: Academic Press. 
PETERSEN, R.S. 1973. The rust fungus life cycle. Bot. Rev. 40:453-513. 
PREECE T.F. & HICK A.J. 1989. An introductory scanning electron microscope atlas of rust fungi. 

London: Farrand Press. 
WILSON, M. & HENDERSON, D.M. 1966. British Rust Fungi. Cambridge: University Press. 



FERN GAZ. 13(6) 1990 329 



SOME ASPECTS OF WATER RELATIONS OF EQUISETUM 
TELMATEIA (EQUISETACEAE: PTERIDOPHYTA) 

RALPH DAVID, UWE PETERS and H. WILFRIED BENNERT 

Spezielle Botanik, Fakultat fur Biologie, Ruhr-Universitat Bochum, 
Universitatsstrasse 150, D-4630 Bochum 1, Federal Republic of Germany 

ABSTRACT 
Diurnal courses of water potential, osmotic potential and pressure potential were studied 
in the Great Horsetail (Equisetum telmateia) growing in a wet alderwood stand (Carici 
remotae-Fraxinetum) in the northern parts of the city of Bochum (North Rhine- Westphalia, 
FRG). The water relation parameters followed in general the characteristic daily pattern 
described for flowering plants. Although the watertable was near the soil surface Equisetum 
telmateia was unable to keep a favourable water balance under certain microclimatological 
conditions (high solar radiation, high vapour pressure deficit) indicating insufficient water 
uptake and/or transport. A marked but temporary nightly decrease of water potential 
was observed repeatedly at various times during a two years' study period, a phenomenon 
for which presently no satisfactory explanation can be given. Methodical problems of 
determining reliable values for osmotic and pressure potential are briefly discussed. 

INTRODUCTION 
Little recent research has been done in the field of water relations of pteridophytes. The 
investigations concentrate on tropical or desert ferns and the influence of droughting on 
the water balance or gas exchange of those plants (Nobel 1978; Nobel et al. 1978; Eickmeier 
1979; Froebe & Strank 1982; Starnecker & Winkler 1982; Sinclair 1983a, 1983b, 1984; Hew 
1984; Winter et al. 1986; Nasrulhaq-Boyce & Haji Mohamed 1987). Some ecophysiologically 
orientated work has been done on the hydraulic conductances and the anatomy of xylary 
elements of different fern species (Woodhouse & Nobel 1982; Calkin et al. 1985; Gibson 
et al. 1984, 1985). 

Detailed data from field measurements focussing on water relation parameters of 
pteridophytes seem to be not available. In the case of the horsetails (Equisetum) this obviously 
reflects the fact that most species represent wetland or swamp plants and that for these 
water supply is not regarded as being critical. There is evidence, however, that plants growing 
in this sort of environment may face specific problems concerning the maintenance of a 
favourable water balance under large air to leaf water vapour concentration gradients; the 
low oxygen concentrations in waterlogged soils may be involved herein (cf. Jones 1971; 
Jones & Muthuri 1984). 

Within a more comprehensive research programme to elucidate the ecophysiological 
adaptations of characteristic species growing in an wet alderwood (Peters 1988) this study- 
concentrates on the daily and seasonal pattern of water relation parameters of Equisetum 
telmateia Ehrh., the Great Horsetail. This species can be a dominant and quite impressive 
plant in certain vegetation types in Central Europe; its contribution to the production and 
mineral cycling within these ecosystems is substantial (Peters, unpublished results). 

MATERIALS AND METHODS 

Study site 

Equisetum telmateia was studied at a site situated in the northern parts of the city of 
Bochum, North Rhine- Westphalia, FRG. On gentle slopes below springs with mainly lateral 
movement of water a small stand of a semi-natural wet alderwood with dominating black 
alder (Alnus glutinosa) is to be found. This plant community can be classified as Carici 
remotae-Fraxinetum W. Koch (cf. Bennert & Kaplan 1983). The soil represents a base- 
rich calcareous gley with a pH above 6.0 and a watertable permanently near the surface. 
Among the herbaceous plants the absolute dominance of Equisetum telmateia is most 



330 FERN GAZETTE: VOLUME 13 PART 6 (1990) 



remarkable. In 1984 about 350,000 shoots of Equisetum telmateia were estimated in the 
whole population covering an area of approximately 5,000 m 2 (Peters & Bennert 1987). 

In 1975 parts of the alderwood was cut down and subsequently a moist meadow as 
a replacement community with several taxa of horsetails including Equisetum telmateia 
developed. To evaluate the influence of the tree canopy on the water relations of Equisetum 
telmateia a comparison was made between plants growing in the alderwood and those 
found in the moist meadow. 

Measurement of water relation parameters 

In 1984 and 1985 monthly investigations of the water relations of Equisetum telmateia 
were carried out during the whole vegetation period. Data of diurnal cycles measured over 
a period of 24 hours obtained in July and August of both years, 1984 and 1985, are selected 
here to demonstrate the variability of these parameters. At this time of the year E. telmateia 
is fully developed, towards autumn the tall plants are often damaged and bent by heavy 
rainfall and storms. 

Water potential (vp w ) measurements were performed in triplicates at intervals of (mostly) 
3 hours with a commercial pressure chamber (PMS Instruments, Corvallis, Oregon). Because 
of the considerable size of the horsetail plants only side branches could be used; these 
were taken from the upper third of the plants. Simultaneously separate samples for 
determination of osmotic potential (in duplicates) were harvested and immediately frozen 
in liquid nitrogen and stored in a deep freezer until further examination. After a short 
thawing period (5 min) sap was expressed from this plant material with a specially constructed 
press (Kreeb 1977) and used for determination of the osmotic potential (vj/ s ) employing 
the cryoscopic method (KNAUER Semi-Micro Osmometer, type M). Each sap sample was 
analysed twice. The values of osmotic potential (^> s ) and water potential (^ w ) were used 
to calculate the corresponding pressure potential ("fp). 

Microclimatological measurements 

Air temperature and relative air humidity were monitored with a hygrothermograph. From 
these data vapour pressure deficit (VPD) was calculated. Total short wave radiation was 
registered with a bimetallic actinograph which was placed on the wet meadow neighbouring 
the forest. 

RESULTS 
Both days in July (25.7.1984 and 24.7.1985) that had been chosen for measurements were 
warm and sunny with maximum temperatures of 22° to 23°C (figs. 1A and 2A). Solar 
radiation reached a maximum value of approximately 4.5 J/cm 2 ■ min on both days (figs. 
IB and 2B). In early afternoon VPD rose up to 13 mbar (figs. 1C and 2C). Starting from 
high predawn values (-1 to -2 bar), the water potential decreased more or less slowly 
during the morning to reach its lowest values of -6 to -7 bar around midday (figs. ID 
and 2D). In the following hours water potential recovered gradually. In July 1984, in spite 
of a relatively high VPD, water potential did not drop below -5.5 bar (fig. ID) whereas 
in 1985 under similar conditions -7.5 bar were reached (fig. 2D). In both years a more 
or less distinct but temporary decrease of water potential during night time was observed. 
The osmotic potential followed generally the course of water potential although its diurnal 
variations were less significant. Especially in July 1985 (fig. 2D) it showed only small changes 
within the range of 1 bar. Pressure potential reached its lowest values around noon; in 
1984 it decreased to only 0.5 bar. 

The 16th of August 1984 (fig. 3) was a mainly sunny and warm day which fell into 
a fairly dry time of year - it had not rained for the previous 1 1 days. In the early afternoon 
values of VPD exceeded 10 mbar. The 26th of August 1985 (fig. 4) was colder with a 
maximum temperature of only 1 6°C and a fairly high degree of air humidity (VPD permanently 



WATER RELATIONS OF EQUISETUM TELMATEIA 331 



under 3 mbar). During day time hours water potentials again reached their minima around 
noon (figs. 3D and 4D); especially low values (-11.5 bar) were recorded in 1984. In 1984 
patterns of variation in osmotic potential paralleled those in water potential; in 1985 it 
showed only minor diurnal fluctuations. In both years the course of pressure potential 
was substantially influenced by that of the water potential. In 1984 a water potential lower 
than the osmotic potential was determined around noon and a negative value of pressure 
potential resulted (fig. 3D). 

Fig. 5 shows results obtained on 26th August 1985 when for comparison additionally 
plants of E. telmateia were measured that were growing on the neighbouring open moist 
meadow. The most obvious difference to the forest plants is to be found in water relation 
parameters around noon and in early afternoon. In the open habitat much lower (more 
negative) values of water potential (-9.5 versus -5 bar) occurred with pressure potential 
approaching zero. In both habitats water potential showed a remarkable nightly decrease 
(at 0200 h) for which no explanation can be deduced from microclimatological data. 

DISCUSSION 

Comparative results of water relation characteristics and their diurnal variations in 
pteridophytes and especially horsetails obtained from field measurements are obviously not 
reported in literature. However, the degree of diurnal cycling as well as the values of water-, 
osmotic-, and pressure potential of Equisetum telmateia lie well in the range determined 
for many herbaceous flowering plants (e.g. Losch & Franz 1974; Richter 1976; Curtis & 
Kincaid 1984; Grimme 1984), especially for those from wetland communities (Richter 1976; 
Larcher 1984). 

Plant water status fluctuates diurnally under variable microclimatological conditions. 
The interactions of environmental factors like temperature, relative air humidity, wind 
velocity, vapour pressure deficit and solar radiation, which change spatially and with time, 
are difficult to establish (Elfving et al. 1972). In many cases the daily course of water 
potential mirrors the diurnal variation of radiation, which, besides VPD, is often regarded 
as having the strongest effect (Huzulak 1977; Pereira & Kozlowski 1978; Grimme 1983). 

The most striking result was the obvious difficulty Equisetum telmateia faced occasionally 
when trying to keep a favourable water balance. This became obvious in a sharp decrease 
of water potential with pressure potential approaching zero (or even being estimated as 
negative) and occurred even under rather moderate environmental conditions, but always 
around noon when air temperature and solar radiation reached their daily maximum. As 
the high watertable of the studied alderwood ecosystem indicates that water was freely 
available to the roots of Equisetum telmateia water shortage in the soil cannot be responsible 
for this phenomenon. Calkin et al. (1985), examining the water conduction of xylem elements 
of different fern species, found in some of them the tracheids partially blocked. Although 
horsetails were not included in these studies, the temporary drop of water- and pressure 
potential in Equisetum telmateia is more likely to be caused by insufficient water conduction 
or uptake rather than reduced water availability. Additional support for this explanation 
can be derived from studies on water relations of certain flowering plants. When their 
roots were subjected to flooding conditions, thereby experiencing anaerobiosis, resistance 
to water flow through roots and stem increased indicating partial occlusion of the xylem 
vessels (Andersen et al. 1984). 

On the other hand water potential recovered fairly quickly in the course of the afternoon 
(see esp. figs. 3 and 5), a process to which also a partial closure of stomata could have 
contributed. Data obtained by Peters (1988) in other years show that Equisetum telmateia 
indeed tends to reduce stomatal opening and transpiration rate early in the afternoon under 
conditions of intensive insolation and high vapour pressure deficit of the air occurring during 
summer months. Midday closure of stomata was observed repeatedly, both, in well watered 



332 FERN GAZETTE: VOLUME 13 PART 6 (1990) 



shrubs and food plants under experimental conditions (Lange & Meyer 1979; Lange et 
al. 1982) and in naturally growing wetland plants (Jones 1971; Jones & Muthuri 1984). 
Whereas Lange & Meyer (1979), interpreting their results obtained in apricot and grapevine, 
regard midday depression of stomatal conductance in plants sufficiently supplied with water 
from an ecological point of view as unnecessary and even disadvantageous, Jones & Muthuri 
(1984) give a quite different explanation. Examining the behaviour of plants of a papyrus 
(Cyperus papyrus) swamp in Africa, they propose that the reduction of water flow into 
the roots at high atmospheric demand is directed at reducing the uptake of toxic ferrous 
iron. These iron compounds as well as other toxic elements (especially Mn 2+ ) are abundant 
under anaerobic and reducing soil conditions of many wetland ecosystems (cf. Crawford 
1982; Wheeler et al. 1985; Janiesch 1986). According to Peters (1988) this applies also to 
the soil of the alderwood stand where the investigated Equisetum plants were growing. 

In all cases a more or less obvious temporary decrease in water potential occurred 
during night, mostly between midnight and 0300 h. Such nightly fluctuations have now 
been observed over a period of two years, occurring rather frequently but irregularly. One 
possible explanation to be considered is that a transitory phase of reopening of stomata 
during night time causes increasing transpiration rates which in turn could result in a drop 
of water potential. However, recent investigations on Equisetum telmateia do no support 
this possibility. Peters (1988) could demonstrate that stomata of Equisetum telmateia often 
are kept open to a certain degree during the whole night and that transpiration rate is 
very low because of high air humidity in the alderwood stand. Neither in stomatal opening 
nor in transpiration rates were observed nightly changes that could explain a decrease of 
water potential in the magnitude of 2 to 3 bar. It is remarkable that also alder (Alnus 
glutinosa) plants from the same habitat exhibited a nightly decrease of water potential, 
however, somewhat less frequent than Equisetum telmateia, but again without any indications 
that changes of stomatal opening and in transpiration rate might be involved (Peters 1988). 
In literature, nightly changes in plant water status is rarely reported or commented on 
(Hinckley 1971; Elfving et al. 1972; Hinckley & Ritchie 1973; Cutler et al. 1977). Rather 
well known for many years has been the fact that stomata sometimes open in the dark, 
a phenomenon usually termed "night opening". It is commonly believed to be related to 
an endogenous rhythm, but it can also be substantially enhanced by higher temperatures 
(e.g. Mansfield 1965; Pemadasa 1977). Occasionally the night opening was interpreted as 
being caused by a shortage of oxygen within the plant (Scarth et al. 1933; Levitt 1976) 
and termed "scotoactive opening". 

Under certain microclimatological conditions a water potential more negative than the 
corresponding osmotic potential was measured and consequently a negative pressure potential 
was calculated. Although the existence of negative turgor is regarded as possible by several 
investigators (e.g. Grieve 1961; Kreeb 1960, 1961; Kappen et al. 1972; Beadle et al. 1978) 
it has never been established with certainty whether - with continuing dehydration - pressure 
potential of cells or tissues remains zero or indeed turns negative. What, however, has 
been demonstrated repeatedly is that negative pressure potentials may result as an artefact 
if the (absolute) value of the osmotic potential is underestimated (cf. Tyree 1976). This 
is normally the case when employing the cryoscopic method as during preparation of sap 
the separation between symplasmatic water (containing solutes) and apoplasmatic water 
(relatively pure water contained in the cell wall) is abolished and both fractions are mixed. 
As a result the cryoscopically measured osmotic potentials are generally too high (their 
absolute values too low). Cutler et al. (1977) report errors up to 45%; Markhart et al. 
(1981), using filter paper as model systems, predict errors bettween 10 and 40%. Therefore 
the negative pressure potential found in Equisetum telmateia must be understood as caused 
by the cryoscopical method employed. But even if the recorded negative pressure potential 
is regarded as unrealistic it indicates that Equisetum suffers from (at least moderate) water 



WATER RELATIONS OF EQUISETUM TELMATEIA 



333 



FIGURE 1: Diurnal course of microclimatological parameters and water relation components of 
Equisetum telmateia (alderwood stand) on 25 July 1984. A: Air temperature (T). B: Total short wave 
radiation (St). C: Vapour pressure deficit (VPD). D: Water relation parameters; A : water potential; 
• : osmotic potential; o : pressure potential. 



[°cj 



25 
20 
15H 
10 
5 



[J /cm • minj 



VPD 



f mbarl 



15 



10 
5H 



T 10 

[bar] 



10 




— i 1 1 1 r 

20 22 2U 2 U 



10 12 U 



B 



~i 1 1 1 1 1 — *• 

16 18 20 22 time of day 
C.E.Khr] 




20 22 2L 2 U 6 8 10 12 U 16 18 20 



t C 



22 ti me 

r c t 

v^.tZ.. i 



of day 
[hr] 



—i 1 r^ i i — i 1 \ 1 1 1 i i 

20 22 2L 2 L 6 8 10 12 U 16 18 20 



22 tj me 
C.E.T. 



of day 
[hr] 





2® ^^ time of day 
CE.T.Ihr] 



334 



FERN GAZETTE: VOLUME 13 PART 6 (1990) 



FIGURE 2: Diurnal course of microclimatological parameters and water relation components of 
Equisetum telmateia (alderwood stand) on 24 July 1985. A: Air temperature (T). B: Total short wave 
radiation (St). C: Vapour pressure deficit (VPD). D: Water relation parametes; A : water potential; 
• : osmotic potential; o : pressure potential. 



fC] 



4 A 



20- 

15 

10 

5 




-i 1 1 1 1 r 



i i r 



T 1 1 1 — ► 



20 22 2k 2 L 6 8 10 12 H 16 18 20 22 time of day 



[j/cm m i n] 



6- 

I 

U- 

2 



B 



C.E.T. I hr. 




t r i r 



t 1 1 i 1 1 1 r 



-i — ► 



VPD 

[mbar] 



20 22 2U 2 4 6 8 10 12 H 16 18 20 22 timGofday 
f C C.E.T. [hr] 



15 

10- 

5 




i i i r 



T I I I r 



20 22 24 2 U 6 8 10 12 14 16 18 20 22 



[bar] 



time of day 
C.E.T. [hr] 




time of day 
C.E.T [hr] 



-5- 



WATER RELATIONS OF EQUISETUM TELMATEIA 



335 



FIGURE 3: Diurnal course of microclimatological parameters and water relation components of 
Equisetum telmateia (alderwood stand) on 16 August 1984. A: Air temperature (T). B: Total short 
wave radiation (St). C: Vapour pressure deficit (VPD). D: Water relation parameters; a : water potential; 
• : osmotic potential; o : pressure potential. 



[OC] 



20 
15- 

10" 
5- 



[j/cm 2 



VPD 
[mbar] 



T 

[bar] 



10- 



5 J 



-5- 



-10 



15- 



10 12 U 16 18 20 22 t|meofdQy 

C.E.T.Ihr] 




time of day 
C.E.T.Ihr 



l 1 1 r 

20 22 2L 2 



10 12 % 16 18 20 22 ti ^ e of day 

C.E.T.fhr] 



20 22 2U 2 



6 8 10 



12 



16 18 



^o" 




22 time of day 

C.E.T.Ihr] 



336 



FERN GAZETTE: VOLUME 13 PART 6 (1990) 



FIGURE 4: Diurnal course of microclimatological parameters and water relation components of 
Equisetum telmateia (alderwood stand) on 26 August 1985. A: Air temperature (T). B: Total short 
wave radiation (St). C: Vapour pressure deficit (VPD). D: Water relation parameters; A : water potential; 
• : osmotic potential; o : pressure potential. 



[°c] 



20- 
15- 

10- 



[j/cm 2 -min] 



VPD 15 

[mbar] 



10 
5H 




20 22 2L 2 I 6 8 10 12 % 16 18 20 22 



I B 



time of day 
CE.T.Ihr] 



^M 




i 1 1 1 1 r 



i 1 1 1 r 



20 22 7L 2 k 6 8 10 12 H 16 18 20 22 timeofday 

CE.T.Ihr] 



T 

[bar] 



10 



-5- 



-10- 



■15- 



20 22 2U 2 

_j i i i_ 



8 10 12 

J l L_ 



time of day 
C.E.T.Ihrl 




16 18 20 22 

_1 I l I L 



->-► 




ti me of day 
C.E.T. [hr] 



WATER RELATIONS OF EQUISETUM TELMATEIA 



337 



FIGURE 5: Diurnal course of microclimatological parameters and water relation components of 
Equisetum telmateia (moist meadow) on 26 August 1985. A: Air temperature (T). B: Total short wave 
radiation (St). C: Vapour pressure deficit (VPD). D: Water relation parameters; : water potential; 
• : osmotic potential; o : pressure potential. 



[•c] 



20- 
15- 

10 



Q/c 



m mi 



VPD 

[mbar] 



T 

[bar] 



n]6 
L 

2 



20 

15 

10 

5 



10 



5- 



— 



-5- 



10 



-15- 




—i 1 1 r 

20 22 2L 2 

B 



10 12 H 16 18 20 2 2 



time of day 
C.E.T. [hr] 




20 22 2U 



-J 1 , 1 1 , 1 1 1 r -w 

8 10 12 H 16 18 20 22 time of day 

CE.T.Ihr] 




20 
D 



22 2L 



10 



12 



16 18 20 



22 



time of day 
C.E.T. [hr] 



20 22 2L 2 U 6 




i 1 r 




18 20 22 f|me of day 
C.E.T. [hr] 



338 FERN GAZETTE: VOLUME 13 PART 6 (1990) 



stress under high atmospheric demand. 

Repeatedly another method, the so-called pressure-volume curve technique (see Tyree 
& Hammel 1972 and Ritchie & Hinckley 1975 for details), was successfully applied to 
different plant species yielding more realistic values of osmotic potential (e.g. Kaplan & 
Gale 1974; Roberts & Knoerr 1977; Cutler et al. 1979; Roberts et al. 1980; Clayton-Greene 
1983). Unfortunately, it could not be used in the case of Equisetum telmateia because of 
the inner cave system of the stems and branches which tends to collapse if high pressure 
is applied. Occasionally values of osmotic potentials obtained by pressure-volume curves 
were used to calculate a correction factor for those determined by cryoscopy (cf. Bennert 
& Mooney 1979). Peters (1988) and David (unpublished results) were able to determine 
such a correction factor for Fraxinus excelsior, Alnus glutinosa and Epilobium hirsutum 
growing in or near the studied alderwood stand. It amounted to 3 to 4 bar depending 
on the species under examination. Provided that a factor of the same magnitude would 
also be applicable to Equisetum telmateia at no time would a negative turgor really occur 
in this species. 



ACKNOWLEDGEMENTS 
We thank Mrs Vera Mukielka, Bochum, for her help in preparing the figures. Parts of 
this work were supported by a grant of the "Minister fur Wissenschaft und Forschung 
des Landes Nordrhein-Westfalen" to H.W.B. 



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FERN GAZ. 13(6) 1990 34 1 



THE CHROMOSOME NUMBER OF ANOGRAMMA LEPTOPHYLLA 
(ADIANTACEAE: PTERIDOPHYTA) FROM EUROPE 

HELGA RASBACH 

23 Datscherstrasse, D-78o4 Glottertal, Federal Republic of Germany 

& 

TADEUS REICHSTEIN 

Inst. f. Organ. Chemie der Universitat, 19 St. Johanns-Ring, 

CH-4o56 Basel, Switzerland 

ABSTRACT 
Meiotic counts in spore mother cells of Anogramma leptophylla are reported for plants 
from seven origins: Madeira (2), France (2), Continental Spain (1), Mallorca (1) and 
Switzerland (1). We invariably found n = 26 n in agreement with the questionable count 
of Tutin in Fabbri (1963), Kurita (1971) and Queiros et al. (1988), (all three for material 
from Europe) and Gibby (1986, material from Madeira). These results are, however, 
in disagreement with Brownlie (1958, n = 29, material from New Zealand), Mickel et 
al. (1966, n = 27 and 29, from Mexico), Baroutsis & Gastony (1978, n = 29, from S. 
Africa), and Mehra & Verma (1960, n = c. 56-57), Verma & Khullar (1965, n = c. 58), 
and Khullar in Manton et al. 1986:130, n = 58 plants from India. On the other hand 
we found n = 29 n for Anogramma chaerophylla (precise origin unknown) in agreement 
with Walker (1966) and with Gastony & Baroutsis (1975). The genus Anogramma is 
obviously cytologically not homogeneous. 

INTRODUCTION 
We became interested in the chromosome number of Anogramma leptophylla (L.) Link 
(1841:1377) when the late Irene Manton wrote to one of us (T.R. in litt. 26th June 1985) 
that in order to complete manuscript on "Cytology of the fern Flora of Madeira" (Manton 
et al. 1986) she urgently needed precise chromosome numbers of Anogramma leptophylla 
and of Notholaena marantae (L.) Desv. subsp. subcordata (Cav.) Kunkel (1969:46). We 
were able to provide one for the Notholaena (Manton et al. 1986: 138) but were too late 
for Anogramma which we raised from spores from Madeira. Fortunately Mary Gibby was 
able to obtain fixings collected in the field, and she found n = 26 n (Manton et al. 1986- 
131, foot note, and Mary Gibby 1986). This agrees with the questionable count of Tutin 
in Fabbri (1963: 323), with the results of Kurita (1971: 41-42), and with the recent count 
of Queiros et al. (1988:124) for material from Europe. It is, however, in contrast with 
counts of other workers who reported n = 29 for material from New Zealand (Brownlie 
(1958); n = 27 and 29 for material from Mexico (Mickel et al. 1966); n = 29 for South 
Africa (Baroutsis & Gastony 1978) and n = 58 for India (Verma & Khullar 1965, see Khullar 
in Manton et al. 1986: 130). These results and counts of other species of Anogramma 
are reviewed by Gastony & Baroutsis (1975); Baroutsis & Gastony (1978); Love et al. 
(1977:142-143); Manton et al. (1986:130-131); and by Gibby (1986). We therefore decided 
to continue raising plants from spores and collecting fixings in the field of different origins, 
particularly Europe, to determine their chromosome numbers. For comparison we included 
a specimen of Anogramma chaerophylla (Desv.) Link (1841:138). 

The chromosomes of Anogramma have proved difficult to study (reviewed by Manton 
et al. 1986:130-131); they are minute, often exhibit stickiness and have a tendency towards 
clumping; the presence of B-chromosomes has occasionally been reported. We encountered 
similar difficulties, but whenever ample, good fixings were available, several well spread 
cells always could be found. We observed neither B-chromosomes nor fragments. The 
precautions which we used for fixings taken in the field, to avoid deterioration during 
subsequent transport, are described below. 



342 



FERN GAZETTE: VOLUME 13 PART 6 (1990) 



Anogramma leptophylla is often described as an annual and sometimes as annual to 
biennial (Dostal in Kramer 1984:113). In our experiments it usually behaved as biennial 
(details see sowing TR-6336, spores from Madeira). Spores sown in December 1985 gave 
ample prothalli which were planted in soil in Febr. 1986. They grew fast, covered the pot 
soon, but died down in May-June. Only in the fall of 1986 did many young sporophytes 
come up on the brownish mat, probably from underground tubercles (Goebel 1877). These 
tubercles must obviously have been formed after fertilisation and before the gametophyt^s 
died down. Mature leaves and fixable sori became available in April 1987. Many other 
sowings made in Jan. 1989 (not further reported here) with spores from Spain, France 
and Italy also died down in summer 1989. In some cases a few gametophytes remained 
green till autumn and through the whole next year, but never produced mature sporophytes 
until a full year had elapsed after sowing. The behaviour of A. leptophylla as annual or 
biennial may depend on growing conditions and season of spore germination, but in our 
experiments it always behaved as biennial. 

MATERIAL AND METHODS 
Spores or fixings collected in the field of Anogramma leptophylla were used from the six 
following origins (registration numbers T.R. are given for the living progeny). 

1. (TR-6336) = R. Viane 3063, (Fig. 1) 11th July 1985, Madeira, roadside between Gaula 
and Aguas Mansas, below the Pico das Eirozes, c. 740 m alt. Together with Adiantum 
reniformeh., Asplenium obovatumVw. subsp. lanceolatum P. Silva, A. onopterisL., Ceterach 
lolegnamense Gibby & Lovis (1989:287), Polystichum setiferum (Forskal) Woynar. 




FIGURE 1: Silhouette of pressed frond of plant TR-6336 = A. leptophylla raised from spores from 
Madeira (pressed 1st June 1987). 



ANOGRAMMA LEPTOPHYLLA CYTOLOGY 



343 



2. (TR-6337) = R. Viane 3050, 10th Juy 1985, Madeira, Curral das Freiras, roadside wall 
falling apart, c. 650 m alt. Together with Asplenium onopteris. 

3. (RAS-608 = fixing coll. in the field). W. Bennert, H. Rasbach & K. Rasbach 15th 
April 1988, Spain, Prov. Cadiz, Miel Valley (W. of Algeceiras), c. 90 m alt. 

4. (R.V.-4512 : fixing coll. in the field). R. Viane 3rd April 1989, Switzerland, Kt. Ticino, 
Valle Maggia, at the left side of the small, old track from Bignasco (442 m) to Madonna 
del Monte (734 m) at c. 550 m alt. under trees. One of the few long known localities 
in Switzerland. 



5. s.n. fixed in the field. W. Bennert s.n. 5th April 1989, France, Dept. Alpes-Mar. Rocher 
de Roquebilliere near Cannes, c. 110 m alt., in open situation between bushes. 

6. s.n. fixed in the field, W. Bennert s.n. 7th April 1989, France, Dept. Var, He de Port- 
Cros (lies d'Hyeres), at c.50 m alt., slope beside a path. 

7. (TR-6995) = J. A. Rosello, s.n. 13th April 1989, Balearic Islands, Mallorca, pres de 
Caimari, dans les crevices des talus ombrages sur calcaire (in litt. 14th April 1988). 

8. (TR-7041) = Anogramma chaerophylla. On 3rd March 1988. Mr R. Schweizer 
(Strengelbach, Switzerland) sent us some living prothalli of this species which he had cultivated 
for some time in his greenhouse. They came up as a weed, in pots with orchids which 
he had received from a commercial dealer and from friends. The precise origin is unknown 
but they came from tropical America, perhaps from Venezuela, Bolivia, Ecuador or Peru. 
For determination of a fertile plant he is grateful to Prof. K.U. Kramer (Z). A photograph 
of a plant cult, in Basel is given in Fig. 2. 




FIGURE 2: Silhouette of pressed frond of plant TR-7041 = A. chaerophylla grown from material 
(gametophytes and small sporophytes) obtained from R. Schweizer (pressed 8th July 1985). 



344 FERN GAZETTE: VOLUME 1 3 PART 6 ( 1 990) 



For fixation juvenile leaves with immature, slightly swollen, whitish sporangia were used. 
If some brownish or blackish points (mature spores) were visible on the lowest pair of 
pinnae, the upper part of the leaves was sometimes still in usable state. A leaf, or part 
of it, was immersed into a solution of 1:3 glacial acetic acid: absolute ethanol, freshly mixed 
in a glass tube with plastic stopper and a small label written with pencil (graphite) included. 
If possible these Fixings were kept at c. 0-4° C in a thermos bottle with ice (in the Field) 
or in the refrigerator at home. After c. 1-5 days the liquid was replaced by fresh mixture 
and the Fixings were stored at c. -15° C in the deep-freezer until they could be examined 
cytologically. If this was not possible or if the Fixings had to be mailed for a journey taking 
more than 3 days, the liquid was replaced by 70% aqueous ethanol. This gave sometimes 
quite acceptable results after being kept protected from light up to 2 months at room 
temperature. 

The technique described by Manton (1950:293-299) was followed for squashing spore 
mother cells in meiosis, staining in aceto carmine and preparing permanent slides. Suitable 
cells were examined and analysed in phase-contrast on an "Olympus" microscope "model 
BH 2 with oil immersion and attachment for phase contrast (by H.R.). 

Plants were raised from spores (by T.R.) on agar medium. The medium given in A.F. 
Dyer (1979:282) was used with slight modification (adding NaCl). The following quantities 
are for one litre: 



0.51 g crist. Magnesium sulphate = Mg S0 4 • 7 H 2 
0.12 g Potassium nitrate = KN0 3 
0.17 g crist. Ferric chloride = FeCl 3 • 6 H 2 
1.44 g crist. Calcium nitrate = Ca(N0 3 ) 2 • H 2 
0.25 g Potassium dihydrogen phosphate = KH 2 P0 4 
0.10 g Sodium chloride = NaCl 
16.00 g Agar powder 

The salts are each dissolved in c. 10 ml clean tap water, added in the given order 
to c. 600 ml tap water, rinsed and Filled up to make one litre. We added trace elements 
(Dopp 1973:3-4; Reichstein et al. 1973:135). No Mycostatin was added. The mixture was 
heated in a round bottle flask (on the steam bath) with occasional stirring for c. one hour. 
The slightly turbid homogeneous solution was poured in c. forty Erlenmeyer flasks 65 mm 
diameter to make a c. 15 mm thick layer. The flasks were covered with tinfoil, sterilized 
in steam (100°C) for twenty minutes and covered tight with "parafilm M" (plastic). They 
can be stored for a year in this condition. If an infection becomes visible, the flask can 
be sterilized again. Prothalli when 2-3 mm high, were pricked out in pots with lime free 
mixture (Rasbach et al. 1983:45). 



PLANTS RAISED FROM SPORES 
We give two examples showing slightly different behaviour of prothalli: 
1. TR-6336, spores from Madeira. Sowing on Agar-medium 7th Dec. 1985; pricking of 
prothalli on soil in one pot 12th Febr. 1986. The prothalli were covered with a plastic 
cup and sprayed daily. They grew well and soon covered the whole surface of the pot. 
They became brown in c. June 1986 and were then kept as before covered in the humid 
greenhouse atmosphere, but not watered. We started to spray again on 1st Nov. 1986 and 
young sporophytes soon started to grow out of the brown mat. Twelve of them were planted 
in single pots kept covered for a few days, then uncovered in the greenhouse (6-20°C) 



ANOGRAMMA LEPTOPHYLLA CYTOLOGY 345 



and sprayed daily. Ten plants survived. Mature leaves and good fixings were available on 
13th Apr. 1987, i.e., 16 months after sowing and after a resting period. Fig. 1 shows two 
fronds of a mature plant. 

2. TR-6995, spores from Mallorca sown 26th Apr. 1988 on Agar-medium gave ample 
prothalli which were pricked out on soil on 19th Sept. 1988. They grew well and most 
decayed again in c. Nov. 1988, but some remained green throughout 1989 and are still 
green (Febr. 1990). The first sporophytes could be potted on 6th July 1989. Good fixings 
were made, fronds were available on 1 1th Jan. 1990. We did not check whether the sporophytes 
came from the green prothalli or from tubercles of decayed ones. 

Most of the sowings (made in Jan. and Apr. 1989, not reported here) behaved like 
TR-6336 i.e. dying down completely in summer, but in a few cases some prothalli remained 
green as TR-6995. Fertile sporophytes were never available within a period of one year 
from sowing. 



ANOGRAMMA CHAEROPHYLLA 
TR-7041. As mentioned, Mr Schweizer (with litt. of 3.3.1988) provided us with a clump 
of small sporophytes (weeds on orchids) which were potted immediately. After eliminating 
some other weeds (Pteris, Dryopteris) we were able to raise 3 specimens of the Anogramma 
(det. by Prof. K.U. Kramer, Z). Mature leaves (Fig. 2) and good fixings were available 
on 3rd Apr. 1989, a whole plant was pressed on 8th July 1989. 



CYTOLOGY 
Five of the seven specimens of A. leptophylla (listed above) gave precise results with n = 26 n 
in meiosis. Specimen TR-6337 (from Madeira) was not examined, because two other specimens 
(including the result of M. Gibby) have already been counted and n = 26 n was found. 
Specimen 5 (W. Bennert s.n. near Cannes) did not yield any good cells and only an approximate 
count n = c. 26 n (± 1-2, surely not 29) could be obtained. Figs. 3-4 give photographs 
and explanatory diagrams for the five mentioned specimens. Fig. 5 gives similar illustrations 
for our material of A. chaerophylla showing n = 29". 

DISCUSSION 

The name Anogramma leptophylla (L.) Link (1841 : 137) is based on Polypodium leptophyllum 
Linnaeus (1753:1092). A lectotype for this name was designated by Morton (1970:101-103) 
and met with general acceptance. It is specimen number 1251156 in LINN. It bears the 
name Polypodium heterophyllum in Linnaeus' hand and the number 46. As Pichi Sermolli 
(1966:496-595) pointed out, Linnaeus called it this before realizing that he had already 
applied the name P. heterophyllum to another species, and published it as P. leptophyllum. 
Linnaeus (1753:1092) gives: "Habitat in Hispania, Lusitania, Galloprovincia". We have 
examined material from all three of these countries and found them to have n = 26 11 . 
We therefore conclude that this is the correct number for A. leptophylla s.str. 

There remains the problem of the plants from New Zealand, S. Africa and India for 
which n = 29 11 has been reported. It would be desirable to check these results. So far, 
we were able to obtain spores from N.India (S.P. Khullar). They gave some prothalli, but 
no sporophytes were produced. We were not able to obtain material from N.Z. and from 
S.Africa. If the reported counts (n = 29") are correct, such material would have to be 
treated as an aneuploid or as a distinct taxon. 



346 FERN GAZETTE: VOLUME 13 PART 6 (1990) 



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FIGURE 3: 3.1, 3.2 and 3.3 = photographs, 3.1', 3.2' and 3.3' = explanatory diagrams of spore 
mother cells in meiosis of A. leptophylla: 3.1 and 3.1' = TR-6336 raised from spores from Madeira, 
in diakinesis showing n = 26"; 3.2 and 3.2' = Ras-608 fixed in the field, Miel Valley, Spain, 
in diakinesis showing n = 26"; 3.3 and 3.3' = RV 4512 fixings collected in the field from 
Switzerland, in metaphase I showing n = 26". 



ANOGRAMMA LEPTOPHYLLA CYTOLOGY 347 









H 
S A* 



4.f 4f 



ill* X2- 

4.2 42' X \ 



5.7 *^ 5/ 



d * ** - - •-" 10 \Jn\ 



FIGURES 4 and 5 (continuation of Fig. 3): 4.1. and 4.1' = A. leptophylla fixings collected in the 
field, W. Bennert s.n. from He de Port-Cros, France, in metaphase I showing n = 26 n ; 4.2 and 
4.2' = TR-6995 = A. leptophylla ex spores from Mallorca, in metaphase I showing n = 
26 n ; 5.1 and 5.1' = A. chaerophylla from Central- or S. America, cell in metaphase I showing 
n = 29". 



348 FERN GAZETTE: VOLUME 13 PART 6 (1990) 



ACKNOWLEDGEMENTS 
We express our gratitude to the following people: 

M. Boudrie (Clermont-Ferrand, France), Dr W. Bennert (Bochum, W. Germany), 
R. Deschatres (Bellerive s/A, France), Prof. S.P. Khullar (Chandigarh, India), D. Marchetti 
(Massa, Italy), R. Prelli (Lamballe, France), J. A. Rossello (Palma de Mallorca, Spain), 
Dr A.E. Salvo Tierra (Malaga, Spain), R. Viane (Brugge, Belgium) for sending us viable 
spores and Fixings; R. Schweizer (Strengelbach, Switzerland) for sending us little living prothalli 
and sporophytes of A. chaerophylla; Dr Mary Gibby (London, G.B.) for information on 
unpublished results (in press) and her help in preparing the manuscript; Prof. K.U. Kramer 
(Zurich, Switzerland) for identification of A. chaerophylla and his help in correcting the 
manuscript. 



REFERENCES 
BAROUTSIS, J.G. & C.J. G ASTON Y 1978. Chromosome numbers in the genus Anogramma II. 

Amer. Fern Journ. 68(1): 3-6. 
BROWNLIE, G. 1958. Chromosome numbers of New Zealand ferns. Trans. Roy. Soc. New Zeal. 

55:213-216. 
DOPP, W. 1927. Untersuchungen iiber die Entwicklung von Prothallien einheimischer Polypodiaceen. 

Pflanzenforschung 8: 3-10. 
DOSTAL, J. in K.U. Kramer (Ed.) 1984. Gustav Hegi. Illustrierte Flora von Mitteleuropa I. 1. 

Pteridophyta. 3. Aufl. Berlin-Hamburg: Paul Parey. 
DYER, A.F. (Ed.) 1979. The experimental biology of ferns. London-New York-San Francisco: Academic 

Press. 
FABBRI, F. 1963. Primo supplemento alle Tavole chromosomiche delle Pteridophyta di Alberto Chiarugi. 

Cariologia 18: 237-335. 
GASTONY, C.J. & J.G. BAROUTSIS 1975. Chromosome numbers in the genus Anogramma. Amer. 

Fern Journ. 65(3): 71-75. 
GIBBY, M. 1986. A chromosome count for Anogramma leptophylla in Madeira. Fern Gaz. 13(2): 

120. 
GIBBY, M. & J.D. LOVIS 1989. New ferns of Madeira. Fern Gaz. 13(5): 285-290. 
GOEBEL, K. 1877. Entwicklungsgeschichte des Prothalliums von Gymnogramme leptophylla Desv. 

Bot.Zeitung 35:61 1-71 
KUNKEL, G. 1969. Sobre Notholaena marantae (Sinopteridaceae). Cuad. Bot. Canar. 5:46-47. 
KURITA, S. 1971. Chromosome study of four species of leptosporangiate ferns Ann. Rep. Foreign 

Students Coll. Chiba Univ. 6: 41-43. 
LINK, H.F. 1841. Filicum species in Horto regio botanico Berolinensis cultae. Berolini. 
LOVE, A., D. LOVE & R.E.G. PICHI SERMOLLI 1977. Cytotaxonomical Atlas of the Pteridophyta. 

J. Cramer FL-9490, Vaduz. 
MANTON, I., J.D. LOVIS, G. VIDA & M. GIBBY 1986. Cytology of the fern flora of Madeira. 

Bull. Br. Mus. nat. hist. (Bot.) 15(2): 123-161. 
MICKEL, J.T., H.W. WAGNER jr. & K. LIM CHEN 1966. Chromosome observations on the ferns 

of Mexico. Caryologia 19(1): 95-102. 
MORTON, C.V. 1970. The lectotype of Polypodium leptophyllum L. Amer. Fern Journ. 60(3): 101- 

103. 
PICHI SERMOLLI, R.E.G. 1966. Adumbratio Florae Aethiopicae 13. Hemionitidaceae. Webbia 21(1): 

487-505. 
QUEIROS, M., J. ORMONDE & J. NOGUEIRO 1988. Notas cariologicas e fitogeograficas de algumas 

pteridophyta de Portugal, I. Acta Bot Malacitana 13: 121-140. 
RASBACH, H., T. REICHSTEIN & J. SCHNELLER 1983. Five further natural hybrids in the genus 

"Cheilanthes" Sw. (Sinopteridaceae, Pteridophyta). Webbia 87(1): 43-62. 
REICHSTEIN, T., J.D. LOVIS, W. GREUTER & J. ZAFFRAN 1973. Die Asplenien der Insel Kreta. 

Ann. Mus. Goulandris 1: 133:163. 
VERMA, S.C. & S.P. KHULLAR 1965. Cytology of some W-Himalayan Adiantaceae sensu Alston 

with cytotaxonomic comments. Cariologia 18: 85-106. 
WALKER, T.G. 1966. A cytotaxonomic survey of the pteridophytes of Jamaica. Trans. Roy. Soc. 

Edinburgh 66: 27-237. 



FERN GAZ. 13(6) 1990 349 



ASPLENIUM x ARTANENSE (ASPLENIACEAE: PTERIDOPHYTA) 
A NEW DIPLOID HYBRID FROM MALLORCA, SPAIN 

J.A. ROSSELLO 

Departament de Biologia i Ciencies de la Salut, Botanica, Universitat de les Hies 

Balears, 07071 Palma de Mallorca, Spain 

P. CUBAS 

Departamento de Biologia Vegetal II, Facultad de Farmacia, 
Universidad Complutense, 28040 Madrid, Spain 

& 

J.L. GRADAILLE and B. SASTRE 
Museu Balear de Ciencies Naturals, Camp d'En Prom, Soller, Mallorca, Spain 

ABSTRACT 

A new Asplenium hybrid from Mallorca (Arta) is here described and named as Asplenium 
x artanense Rossello, Cubas, Gradaille & Sastre. This hybrid is diploid showing up to 
72 univalents at meiosis. The cytological results, as well as the morphological characteristics 
and ecology of this plant, strongly suggest that it originated from a cross between A. 
sagittatum and A. trichomanes subsp. inexpectans. 

INTRODUCTION 
The European species of Aspleniaceae have been ascribed to either five different genera: 
Asplenium L., Phyllitis Hill, Ceterach DC, Phyllitopsis Reichstein and Pleurosorus Fee 
(Pichi Sermolli 1977, 1987; Reichstein 1981; Ferrarini et al. 1986) or to a single genus: 
Asplenium (Jermy 1968; Lovis & Vida 1969; Salvo et al. 1982; Derrick et al. 1987). Accordingly 
with the adopted taxonomic position, the natural hybrids between species of Phyllitis and 
Asplenium have been named either as intergeneric (x Asplenophyllitis sp.) or as intrageneric 
hybrids (Asplenium x sp.). Since the existence of these hybrids demonstrates that these 
two groups are not completely genetically isolated (Lovis & Vida 1989), we consider that 
there are no strong reasons for splitting Asplenium into different genera, and therefore 
all the European taxa of Aspleniaceae are here treated as belonging to a single genus: 
Asplenium. 

A. scolopendrium L. and A. sagittatum (DC.) A.J. Bange are diploid species (Manton 
1950), and probably both of them are descendants from a common ancestor (Emmott 1964). 
A. scolopendrium is known to have crossed with A. trichomanes L. subsp. quadrivalens 
D.E. Meyer, A. adiantum-nigrum L., A. obovatum Viv. subsp. lanceolatum P. Silva and 
A. lepidum C. Presl subsp. lepidum, thus originating four wild triploid hybrids named: 
A. x confluens, A. x jacksonii, A. x microdon and A. kummerlei (Vida 1963; Girard & 
Lovis 1968; Lovis & Vida 1969; Lovis 1975). 

The related species A. sagittatum is involved in the origin of A. hybridum (Milde) A.J. 
Bange (= Phyllitopsis hybrida (Milde) Reichstein), an allotetraploid species whose parentage 
is A. sagittatum and A. ceterach L. subsp. bivalens (D.E. Meyer) Greuter & Burdet (Vida 
1963; Emmott 1964). Apart from this fertile species, only one hybrid involving A. sagittatum 
as one of the parents has been described till now in Europe: A. x dutartrei. Based on 
its morphological characteristics, this hybrid has been considered to be the cross between 
A. sagittatum and A. ceterach subsp. ceterach (Berthet 1981). 

During field work in the Balearic Islands, in 1987, a plant was found in Mallorca whose 
morphology, ecology and cytology strongly suggest that we are dealing with a hybrid involving 
either A. sagittatum or A. scolopendrium, and A. trichomanes. The description of this 



350 



FERN GAZETTE: VOLUME 13 PART 6 (1990) 



plant as a new hybrid named A. x artanense as well as a discussion on its proposed parents 
follow. 

MATERIAL AND METHODS 
Only one plant of this hybrid has been found in Mallorca, therefore a decision in order 
to keep it alive in the field was made. Only a limited number of fronds were cut to be 
pressed or fixed, and to serve as herbarium specimen. The description of the morphology 
is based on field notes and pressed fronds. Developing sporangia were fixed in the field, 
stained, squashed and preparations made permanent according to the method of Manton 
(1950). 






FIGURE 1: Drawing of the lower side of a frond of A. x artanense. 




1cm 



MORPHOLOGY OF THE HYBRID 
Fronds up to 6 cm x 1.5 cm, narrowly triangular with undulate margins (Fig. 1). Petiole 
about 1/3 as long as lamina, sparsely covered with scales. Lamina coriaceous, dark green 
in the upper side and light green in the lower side, with sparse scales; pinnatipartite for 
the lower 2/3 of its length, the upper third being entire. Basal pinnae almost orbicular, 
base cordate or cuneate, crenate. Middle pinnae broadly elliptic, crenate. Upper pinnae 
subrectangular, ill-defined (indistinct). Rachis brown in the lower half, green to the apex, 
with numerous scales. Sori divergent, 5-8 mm, single or in pairs. Sporangial content consisting 
of nearly misshapen spores of variable size (Figs. 2 A-D). Scales 0.4-1 mm, entire, subulate, 
dark brown without a central occluded area (Fig. 2 E). 

This plant shows an intermediate morphology between A. trichomanes and either A. 
sagittatum or A. scolopendrium (Fig. 3). The texture of the lamina, the presence of some 
sori grouped in pairs, and the morphology and abundance of scales on the petiole, rachis 
and lamina indicate the influence either of A. sagittatum or A. scolopendrium. However, 
the outline of the lamina, slightly wider at the base than at the apex, suggests the influence 
of the cordate or hastate base of A. sagittatum. The degree of dissection of the lamina, 
the crenulate margin and the divergent sori show the characters of A. trichomanes. 



ASPLENIUM x ARTANENSE HYBR.NOV. 



351 



FIGURE 2: Photomicrographs of spores and scales of A. x artanense. A: Sporangial content with 
mainly mi&shapen spores; B: Detail of some shrunken and ill-developed spores; C and D: S.E.M. 
pictures of a spore and detail of perispore pattern; E: Rachis and petiole scales. 





0.5 mm 






352 



FERN GAZETTE: VOLUME 13 PART 6 (1990) 



FIGURE 3: Silhouettes of A sagittatum (A), A. x artanense (B and C), and A. trichomanes (D) 
from Arta, Puig d'En Xiroi. 








CYTOLOGY OF THE HYBRID 

This plant is a diploid hybrid showing an irregular meiosis (Fig. 4), with up to 72 unpaired 
chromosomes at metaphase I. In some cells 1 bivalent was seen, together with 70 univalents. 
Later stages of the meiosis are irregular resulting in the formation of nuclei of different 
size, and univalents lost in the cytoplasm. About half of the chromosomes are larger than 
the others. These results indicate that two different genomes, showing no synaptic homology 
in meiosis, are present in this plant. 

ECOLOGY OF THE HYBRID 
The hybrid plant grows in a rock crevice, at the base of the NW-facing steep-walled outcrop 
of carbonate rocks. Other plants restricted to this type of environment were found in the 
vicinity: Galium crespianum J. Rodr., Digitalis minor L., Micromeria filiformis (Ait.) Benth., 
Crepis triasii (Camb.) Nyman, and Sesleria insulahs Sommier. Several plants of A. 
trichomanes and A. sagittatum were also found in the immediate vicinity (Fig. 3). 



DISCUSSION 
This diploid hybrid must have originated from the cross of two diploid plants with no 
homology between their genomes. The morphology strongly supports that the hybrid is 
a cross between a diploid A. trichomanes with either A. sagittatum or A. scolopendrium. 
Besides, distinct differences in size of the individual chromosomes present in the hybrid 
support this hypothesis. Manton (1950) shows that A. scolopendrium presents larger 
chromosomes than other Asplenium species, a result confirmed in wild and synthetic hybrids 
involving A. scolopendrium as one parent (Girard & Lovis 1968; Lovis & Vida 1969; Vida 



ASPLENIUM x ARTANENSE HYBR.NOV. 353 



FIGURE 4: Cytology of A. x artanense. A: Metaphase I, 1" + 70 1 ; B: Metaphase I, 72' ; C: Anaphase 
II with lagging chromosomes, some appear to be splitting; D: Telophase II, three nuclei of different 
size and a few chromosomes. 



A 


f»M 




f 




f 
• 






*# 




% 


• 




% 










<* 




% 


* 


* % » 


« 






% 


• * ' 


| 


1 f 




• 


* 






1 


-- 




1 * 








,/'• 






| 







B 



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D 



M 



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IQ jjrr 



1970). A. sagittatum seems to have chromosomes of similar size. to those of A. scolopendrium 
(Emmott 1964, PL 22-8). 

Both A. sagittatum and A. scolopendrium grow in Mallorca, however, they are associated 
with different habitats in the island. A. scolopendrium grows either within vertical cavities 
in a mostly karstic landscape or within crevices in the narrow and steep-walled bounding 
sides of small streams (locally known as "torrents"). In both cases, shade and high humidity 
characterize the sites in which this plant grows. A. sagittatum endures drier and brighter 
conditions. In the Arta region (Eastern Hills) only A. sagittatum has been found and, 
in fact, is growing nearby the hybrid. Thus, the ecology of the hybrid site as well as the 
presence of A. sagittatum in the vicinity of the hybrid strongly suggest that A. sagittatum 
is one of the parents of the hybrid. Therefore, the large chromosomes observed in the 
meiosis of the hybrid must have been supplied by the A. sagittatum parent. 

Concerning the other parent of the hybrid, another unrelated diploid taxon has to be 
chosen in order to explain the cytological behaviour of the plant. Again, the morphology 
and field spatial relationships between the hybrid and A. trichomanes plants, lead us to 
consider this taxon as the other parent of the hybrid. Following the taxonomy proposed 
for this aggregate species by Lovis (1964) and Lovis & Reichstein (1985), diploid plants 



354 FERN GAZETTE: VOLUME 13 PART 6 (1990) 



of A. trichomanes growing on calcareous rocks may be included in the subsp. inexpectans. 
However, it should be noted that some of the A. trichomanes plants growing in the Arta 
region, deviate from the morphology of subsp. inexpectans. These plants have small spores 
(mean exospore length ranging from 29.5 to 30.2 urn), thus indicating that they probably 
are diploid. However, they have narrow pinnae with serratolobate margins, a feature which 
recalls the morphology of the tetraploid subsp. pachyrachis, a taxon not found in Mallorca. 
Moreover, plants from other places of Mallorca, with similar morphology and mean exospore 
length, have been cytologically checked, and have proved to be diploids (e.g. samples from 
Mortitx, mean spore length = 27.27 urn, n = 36 bivalents). 

Further research is in progress to elucidate whether: a) subsp. inexpectans is a highly 
polymorphic taxon, and therefore, in spite of their morphological variability, all the diploid 
plants growing in Mallorca can be ascribed to this subspecies, or b) another diploid taxon 
within the A. trichomanes complex (apart from subsp. inexpectans and subsp. trichomanes, 
and closely related to subsp. pachyrachis) should be considered, and deserve a formal 
taxonomic recognition. 

Since this topic may remain as an open question for some time, we ascribe all the 
diploid A. trichomanes plants found on carbonate rocks in Mallorca to the subsp. inexpectans 
and so we propose that most probably a plant of this subspecies was involved in the origin 
of the hybrid plant. 

CONCLUSION 
Based on the above presented evidences, we conclude that the studied diploid hybrid has 
resulted from the cross of A. sagittatum and A. trichomanes subsp. inexpectans, and we 
propose to name this new hybrid as A. x artanense, whose formal description follows: 

Asplenium x artanense Rossello, Cubas, Gradaille and Sastre, hybr. nov. (= A. sagittatum 
(DC.) Bange x A. trichomanes L. subsp. inexpectans Lovis). 

Diagnosis: Planta hybrida diploidea, media inter parentes. Frondes usque ad 6 cm longa 
et 1.5 cm lata. Fronde angustate triangulare, pinnatipartita, paleacea, basi auriculata. Sporae 
abortivae. Meiosi chromosomatibus univalentibus 70-72 et bivalentibus 0-1. 

Holotypus (Fig. 5): Spain, Mallorca, Arta, Puig d'En Xiroi, 15.5.1989, 200 m alt., in 
a limestone crevice, J.L. Gradaille et al., MAF 131406, growing with A. sagittatum and 
A. trichomanes subsp. inexpectans. 

Derivatio: Named after the town of Arta (Mallorca). 

FIGURE 5: Fronds of A. x artanense (holotypus). 




1 2 CM 

1 ■' ! 1 1 1 ' 1 1 1 1 " 1 1 1 



ASPLENIUM x ARTANENSE HYBR.NOV. 355 



REFERENCES 
BERTHET, P. 1981. Un nouvel hybrid d' Asplenium: A. dutartrei (Fougeres). Bull. MensuelSoc. Linneene 

de Lyon 50(8): 250-253. 
DERRICK, L.N., JERMY, A.C. & PAUL, A.M. 1987. Checklist of European pteridophytes. 

Sommerfeltia 6: 1-194. 
EMMOTT, J.I. 1964. A cytotaxonomical investigation in the Phyllitis- Asplenium complex. New 

Phytologist 63: 306-318. 
FERRARINI, E., CIAMPOLINI, F., PICHI SERMOLLI, R.E.G. & MARCHETTI, D. 1986. 

Iconographia palynologica pteridophytorum Italiae. Webbia 40: 1-202. 
GIRARD, P.J. & LOVIS, J.D. 1968. The rediscovery of x Asplenophyllitis microdon, with a report 

on its cytogenetics. Brit. Fern Gaz. 10: 1-8. 
JERMY, A.C. 1968. Review of "Excursion Flora of the British Isles". Brit. Fern Gaz. 10: 51. 
LOVIS, J.D. 1964. The taxonomy of Asplenium trichomanes in Europe. Brit. Fern Gaz. 9: 147-160. 
LOVIS, J.D. 1975. Asplenium L. x Phyllitis Hill = Asplenophyllitis Alston, pp. 104-106. In Stace, 

C.A. (Ed.). Hybridization and the flora of the British Isles: London, New York, San Francisco: 

Acad. Press. 
LOVIS, J.D. & VIDA, G. 1969. The resynthesis and cytogenetic investigation of x Asplenophyllitis 

microdon and x A. jacksonii. Brit. Fern Gaz. 10: 53-67. 
LOVIS, J.D. & REICHSTEIN, T. 1985. Asplenium trichomanes subsp. pachyrachis (Aspleniaceae, 

Pteridophyta), and a note on the typification of A. trichomanes. Willdenovia 15: 187-201. 
MANTON, I. 1950. Problems of cytology and evolution in the Pteridophyta. Cambridge: Univeristy 

Press. 
PICHI SERMOLLI,, R.E.G. 1977. Tentamen pteridophytorum genera in taxonomicum ordinem 

redigendi. Webbia 31: 313-352. 
PICHI SERMOLLI, R.E.G. 1987. A look at the chromosome numbers in the families of Pteridophyta. 

Webbia 41(2): 305-314. 
REICHSTEIN, T. 1981. Hybrids in European Aspleniaceae (Pteridophyta). Bot. Helv. 91: 89-139. 
SALVO, A.E., PRADA, C. & DIAZ, T. 1982. Revision del genero Asplenium L., subgenero Pleurosorus 

(Fee) Salvo,, Prada & Diaz. Candollea 37: 457-484. 
VIDA, G. 1963. A new Asplenium (sectio Ceterach) species and the problem of the origin of Phyllitis 

hybrida (Milde) Christ. Acta Bot. Acad. Sci. Hung. 9: 197-215. 



356 FERN GAZETTE: VOLUME 13 PART 6 (1990) 

THE FERN GAZETTE 



NOTES FOR CONTRIBUTORS 

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Diagrams and photographs should be in black-and-white, of the required magnification 
or larger, with identity and top edge indicated. 



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FERN GAZ. 13(6) 1990 357 



A NEW NATURAL HYBRID IN THE GENUS PTERIS (PTERIDACEAE: 
PTERIDOPHYTA) FROM THE KUMAUN HIMALAYA 

Y.P.S. PANGTEY AND S.S. SAMANT 

Botany Department, D.B.S. Campus, 

Kumaun University, Nainital, India 

& 

SANDEEP VERMA 
Botany Department, Panjab University, Chandigarh, India 

ABSTRACT 

Pteris x khullari a new natural hybrid is described. One of the parents is presumably 
Pteris wallichiana. This is the only report of a natural hybrid of the genus Pteris from 
the W. Himalaya. 

INTRODUCTION 

In the recent past, natural hybrids have been widely assumed to be absent in the Himalaya, 
as the species were mostly not properly known. Hybrids can be detected by the presence 
of a high percentage of abortive spores, combined with an unusual morphology intermediate 
between the parents, and also irregular meiosis. 

Till recently, only a single natural hybrid from the W. Himalaya was known (a diploid 
hybrid Athyrium x pectinatum Mehra & Bir 1960). Later Fraser- Jenkins (1986) added 5 
more natural hybrids of the genus Dryopteris and 2 in the genus Polystichum (pers. common, 
to S.P. Khullar). Two natural hybrids in the genus Cystopteris were reported by Khullar 
(1983) from the Patnitop area (Kashmir). In addition to these, the common natural hybrid 
Asplenium x alternifolium was known from Kashmir (Stewart 1945, 1972), the latter also 
found in Kinnaur, Himachal Pradesh (Khullar & Sharma 1987). 

THE DISCOVERY 

During a survey of the ferns of the Kumaun Himalaya, an unusual taxon was collected 
from Pithoragarh district. On examination it was found to be a hybrid with a high percentage 
of abortive spores. This taxon belongs to the Section Campteria of the genus Pteris, as 
it possesses the typical costular areolae. To this section belong only 2 species in the W. 
Himalaya viz: Pteris biaurita L. (diploid apomictic and triploid apomictic Manton & Sledge 
1954, Verma in Mehra 1961) and Pteris wallichiana Ag. (diploid sexual, Love et al. 1977). 
One of the likely parents of the present taxon is P. wallichiana. 

This taxon has been named after Dr S.P. Khullar, Botany Department, Panjab University, 
Chandigarh (India) in recognition of his contributions to the study of W. Himalaya ferns 
and also for detecting this hybrid in the fern collections of Kumaun sent to him for 
identifications. 

DESCRIPTION 
Pteris x khullarii Pangtey, Samant et Verma hybrida nova 

Rhizoma erectum, apice squamis fuscis concoloris vestitum. Stipes c.30 cm longus, 
stramineus, crassus, 0.2 cm in diametro, glaber, nitidus. Rhachis stipiti similis, glabra. 
Lamina pinnata vel bipinnata, c.30 cm longa, 20 cm lata, triangulari-lanceolata, 
herbacea, glabra. Pinnae multijugatae, usque ad 15 cm longae, c.10 cm latae; pinnis 
basalibus maximis, alternatis, petiolatis, pinnulis bene evolutis instructis; pinnis 
distalioribus gradatim minus lobatis demum elobatis; omnibus pinnis margine varie 
lobatis in lobis anguste-linearibus integre aut lobate marginatis incisis. Pinnulae 
multijugatae, c.4-6 cm longae et 0.25-0.3 cm latae, alternatae, breviter petiolatae, in 



358 



FERN GAZETTE: VOLUME 13 PART 6 (1990) 







FIGURE 1: Diagrams of Pteris x khullarii 

a. part of the frond 

b. an ultimate lobe 

c. part of the frond to show venation 

d. various types of pinnules 

e. spores 

parte distale pinnae sensim sessiles, margine integre aut varie lobatae, apice acutae, 

venis liberis, simplicibus, in lobis furcatae sed venis binatis basalibus acroscopicis 

basiscopicisque anastomosantibus series areolarum costulatarum formantibusque. Sori 

marginales indusiati lobis varie fertilibus. Sporae fuscae, pro parte maxima abortivae. 

Holotypus: W. Himalaya, Uttar Pradesh, Kumaun, Maupani, between Thai and Didihat 

about 0.5 km from Sandeo'by the short route towards Thai, exposed place in the forest, 

c.2000 m alt., coll. Pangtey & Samant SPK 70. 

Rhizome erect, apex scaly, scales brown, concolorous. Stipes c. 30 cm long, stramineous, 
thick, 0.2 cm dia., glabrous, glossy. Rhachis similar to stipe, glabrous. Lamina 1- 
2 pinnate, c. 30 cm long, 20 cm broad, triangular-lanceolate, herbaceous, glabrous. 
Pinnae many pairs, up to 15 cm long, c. 10 cm broad. Basal pinnae the largest, 



PTERIS x KHULLARII HYBR.NOV. 



359 







FIGURE 2: Silhouette of Pteris x khullarii 



360 FERN GAZETTE: VOLUME 1 3 PART 6 ( 1 990) 



alternate, petiolate. Pinnae in the distal region of the frond gradually less lobed and 
then lamina only once pinnate. Pinna margins variously lobed into narrow linear 
lobes with entire or lobed margins. Pinnules in many pairs, c. 4-6 cm long, 0.25- 
0.3 cm broad, alternate, shortly petiolate becoming sessile in the distal region of 
the pinna, linear, margin entire or variously lobed, apex acute. Veins free, simple, 
forked in the lobes, but the basal pair of acroscopic and basiscopic veins anastomosing 
to form a row of costular areolae. Sori marginal, indusiate, the lobes variously fertile. 
Spores brown, a large percentage of aborted spores present. 

ACKNOWLEDGEMENTS 
We are highly grateful to Prof K.U. Kramer (Zurich, Switzerland) for confirming the hybrid 
nature of the present taxon. We also wish to express our grateful thanks to Dr Paolo 
Luzzi (Curator, Botanical Gardens, Florence, Italy) for the Latin diagnosis. 

REFERENCES 
FRASER-JENKINS, C.R. 1986. A classification of the genus Dryopteris (Pteridophyta: Dryopteridaceae). 

Bull. Brit. Mus. (Nat. Hist.) London 14(3): 183-218. 
KHULLAR, S.P. 1983. Natural hybrids in Cystopteris and the taxonomy of C. fragilis species complex. 

National Seminar on "Progress in Botanical Research". Eds. S.C. Verma and T.S. Sareen, Dept. 

of Botany, Panjab University, Chandigarh: 82-83. 
KHULLAR, S.P. & SHARMA, S.S. 1987. The ferns of Western Himalaya. In 'Western Himalaya' 

1: 310-346, eds. Y.P.S. Pangtey and S.C. Joshi. Gyanodya Press, Nainital (India) . 
LOVE, A., LOVE, D. & PICHI SERMOLLI, R.E.G. (1977). Cytotaxonomical atlas of the Pteridophyta. 

Vaduz: Cramer. 
MANTON, I. & SLEDGE, W.A. 1954. Observations on the cytology and taxonomy of the pteridophyte 

flora of Ceylon. Phil. Trans. R. Soc. London, B, 238: 127-185. 
MEHRA, P.N. 1961. Chromosome numbers in Himalayan ferns. Res. Bull. Panjab Univ., N.S. 12: 

139-169. 
MEHRA, P.N. & BIR, S.S. 1960. Cytological observations on the Himalayan species of Athyrium 

and comments on the evolutionary status of the genus. Amer. Fern J. 50: 276-295. 
STEWART, R.R. 1945. The ferns of Kashmir. Bull. Torrey Bot. Club 72: 399-426. 
STEWART, R.R. 1972. An annotated catalogue of the vascular plants of West Pakistan and Kashmir. 

In E. Nasir & S.I. Ali (eds), Flora of West Pakistan. Islamabad. 



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The British Pteridological Society 
THE FERN GAZETTE 

VOLUME 13 PART 6 1990 

CONTENTS 



Page 

MAIN ARTICLES 

Scanning electron microscope photographs of the sori and spores of six species 

of Rust Fungi (Uredinales)iound on ferns in Britain 

- Adrian J. Hick & Thomas F. Preece 321 

Some aspects of water relations of Equisetum telmateia (Equisetaceae: Pteridophyta) 

- Ralph David, Uwe Peters & H. Wilfried Bennert 329 

The chromosome number of Anogramma leptophylla (Adiantaceae: Pteridophyta) 
from Europe 

- Helga Rasbach & Tadeus Reichstein 341 

Asplenium x artanense (Aspleniaceae: Pteridophyta) 
A new diploid hybrid from Mallorca, Spain 

- J. A. Rossello, P. Cubas, J.L Gradaille & B. Sastre 349 

A new natural hybrid in the genus Pteris (Pteridaceae: Pteridophyta) 
from the Kumaun Himalaya 

- Y.P.S. Pangtey, S.S. Samant & S. Verma 357 

NOTES FOR CONTRIBUTORS, ETC. 356 



(THE FERN GAZETTE Volume 13 Part 5 was published on 22nd November 1989) 



Published by THE BRITISH PTERIDOLOGICAL SOCIETY, c/o Department of Botany, 
The Natural History Museum, London SW7 5BD 

ISSN 0308—0838 

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FERN GAZ. 13(7) 1990 361 



MODULAR GROWTH OF HUPERZIA SELAGO (LYCOPODIACEAE: 

PTERIDOPHYTA) 

ALISTAIR D. HEADLEY 

Department of Environmental Science, The University, Bradford, West Yorkshire, 

BD7 1DP, England 

& 
TERRY V. CALLAGHAN 

Institute of Terrestrial Ecology, Merlewood Research Station, Grange-over-Sands, 

Cumbria LA 11 6JU, England 

ABSTRACT 
Huperzia selago is an evergreen perennial of arctic and alpine environments. It demonstrates 
modular growth as it consists of a series of annually produced segments which can be 
aged. 

H. selago demonstrates a very deterministic* pattern of growth with early dichotomous 
branching followed by infrequent branching as the lower parts of branches become 
decumbent. Relative growth rates are very low and characteristic of species of stressed 
environments in which it is found. 

H. selago has a high vegetative reproductive capacity through the production of bulbils, 
which are readily established in relatively closed vegetation. The bulbils are more successful 
as a means of vegetative spread in extremely patchy environments than the stononiferous 
habit. This form of reproduction is opportunistic in relatively closed vegetation compared 
to sexual reproduction. The species relies on spore production for long distance transport 
and opportunistic establishment in early successional habitats generated by disturbance 
and erosion on the tops and steep slopes of mountains. 

INTRODUCTION 

The sporophyte of Huperzia selago (L.) Bernh. ex Schrank is a widespread perennial of 
the stressed environments of arctic and alpine areas. It shows an annual cycle in which 
sporophyll and microphyll production by the apical meristem alternate (Case 1943). As 
a consequence, all segments of the plant can be aged (Headley 1986). This means that 
the modular concept (Prevost 1978) can be used to describe the age-related growth, fecundity, 
death and survival of branches (Callaghan et al 1986a; 1986b; 1990). 

The application of the modular concept to the growth and physiology of the closely 
related stoloniferous plant, Lycopodium anontinum L., has been used to describe its foraging 
habit and response to spatially patchy environments (Callaghan et al 1986a). Also it has 
been shown that this species has potentially indefinite growth (Callaghan et al 1990). However, 
H. selago is a decumbent perennial that reproduces vegetatively by bulbils and despite 
having potentially indefinite growth, it is a short-lived perennial with a very different growth 
and reproductive strategy from that of L. annotinum. 

This paper described in detail the modular growth of H. selago growing in three contrasting 
environments, one a temperate upland oceanic site in Snowdonia, North Wales, another 
an alpine subarctic site in northern Sweden and a third high arctic site in Svalbard. 

Sites MATERIALS AND METHODS 

Material was collected from three sites: one below Carnedd Dafydd in North Wales 
(53° 09' N, 3° 58' W,); one on Mount Njulla in the Abisko National Park in Swedish 
Lapland (68° 24'N, 18° 42'E,) and one on Svalbard (78° 15' N, 16° 30' E). The site in 
Snowdonia is an acid grassland at 580 m a.s.l. on a stabilised talus slope. The vegetation 
is dominated by Nardus stricta L. and Festuca ovina L. with Vaccinium myrtillus L., Galium 
saxatile L., Anthoxanthum odoratum L., Cryptogramma crispa (L.) R.Br, ex Hooker, 
Campylopus paradoxus Wils., Hypnum cupressiforme Hedw., Polytrichum alpinum Hedw. 



*n.b. The terms deterministic and opportunistic are used in the sense of plasticity in growth and 
reproductive strategies of the plant, and not used in a morphogenetic sense. 



362 



FERN GAZETTE VOLUME 13 PART 7 (1990) 



and Rhytidiadelphus loreus (Hedw.) Warnst, also present. The site at Abisko is close to 
the summit of Mount Njulla (1 160 m a.s.l.) which supports a dry mountain heath vegetation. 
This is dominated by Salix herbacea L. and Cassiope tetragona (L.) D. Don, with Festuca 
ovina, Vaccinium vitis-idaea L., V. myrtillus, Salix polaris Wg., Silene acaulis (L.) Jacq., 
Carex bigelowii Torr. ex Schwein. and Polytrichum alpinum also present in the vegetation 
(see Headley 1986 for further details). The site on Svalbard is on a south-facing snow- 
free ridge at 200-250 m a.s.l. and the vegetation is dominated by Salix polaris, Cassiope 
tetragona, Polygonum viviparum L. and Carex rupestris All. 

Material 

Branches of H. selago only grow apically and branch by true iso-dichotomy (011gaard 
1979). Senescence proceeds distally with the microphylls and sporophylls remaining in situ. 
There is an annual alternation in the production of sporophylls and microphylls. Sporophylls 
are initated at the beginning of the growing season and the apical meristem changes to 
initiating microphylls at the end of the growing season (Case 1943). The sporangia require 
over a year to mature, but they do not normally dehisce and release the spores until the 
winter after they have matured. Bulbils are generally initiated with the microphylls and 
are supported by specialised structures (bulbil bases) which persist like the microphylls. 
The bulbils and microphylls take a year to mature and bulbils are released in the autumn 
and winter following initiation (Case 1943). This pattern of growth means that annual segments 
of plant can be aged by counting back sequences of sporophylls and microphylls from 
an active meristem (Fig. 1). 



Mature Bulbil 



Mature Sporangi 
Sporophylls 



Sterile 
Microphylls 



Dehisced 
Sporangia 



Bulbil Base 




Current year's 
segment 



, 1-year old 
segment 



> 2-year old 
segment 



^3-year old 
segment 



FIGURE l. A branch of Huperzia selago showing four annual segments of growth. 



MODULAR GROWTH OF HUPERZIA SELAGO 



363 



Over the first 2 to 4 years of the life of a H. selago plant, bulbils and sporophylls 
are not produced. The segments of annual growth can still be identified however, by the 
slight reduction in the size of microphylls produced at the end of the growing season. 
The development of a plant of H. selago therefore enables the determination of overall 
plant age, as well as the ages of all segments, and the sequence of production of all segments 
within the plant (Fig. 2). 

Roots are initiated just behind the apicalmeristem and grow down through the cortex 
(Saxelby 1908). They emerge from the base of the branches where they contact the soil 
and as a consequence they cannot be aged in contrast to the root system of Lycopodium 
annotinum (Headley ef al. 1985). 



SEGMENT 

•fi (yr.) OOSilior 

7 




FIGURE 2. Schematic diagram of two typical Huperzia selago plants (one 7 years old and the other 
4 years old) with the sequence of segment ages and segement position shown. 



Methods 

On 25th March 1982, 19 plants of H. selago were collected from the Snowdonia site and 
stored at -15°C until they were analysed. The plants were divided into their annual segments 
and classified according to their age and position (see Fig. 2). The number of bulbil bases 
on each segment was counted and the number of dichotomies was also recorded. The segments 
were dried at 105°C for 12 hours before being weighed individually. On 2nd August 1982, 
18 plants of H. selago were collected from the Abisko site. They were dried and pressed 
before being analysed as above. Thirty seven plants were collected at random from 
Adventdalen, Svalbard in July 1987 and stored at -20°C prior to analysis. 

A further 20 plants of H. selago were collected from the Snowdonia site complete with 
the soil in which they were rooted. The soil was carefully washed from their roots. The 
plants were aged and separated into roots and branches, before being dried and weighed. 
Ten fresh plants of H. selago were collected from the Snowdonia site on 10th October 
1982 and the mature undehisced sporangia carefully removed and counted from each of 
35 one year old segments. The one-year-old segments, spores and sporangial wall material 



364 



FERN GAZETTE VOLUME 13 PART 7 (1990) 



were dried and weighed as above. The number of spores per sporangia was estimated by 
suspending the spores dehisced from a known number of sporangia in 1 cm 3 water containing 
a wetting agent and counting the number of spores in 0.1 mm 3 of the suspension using 
a haemocytometer slide. Four separate counts were made for each of 3 separate suspensions, 
each containing the spores from 38 or more sporangia. 



75 


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Segment position 



10 11 12 13 



FIGURE 3. Changes in branch fecundity with increasing distance from the base of Huperzia selago 
plants, i.e. segment position, from Snowdonia (circles), Abisko (triangles) and Svalbard (squares). Means 
with S.E. bars shown. 

The viability of spores from fresh branches was estimated by opening a fresh mature 
sporangium in a drop of lactophenol cotton blue (cytoplasmic stain) on a microscope slide. 
The numbers of stained and unstained spores were counted for transects across the slide 
until a total of 600 or more spores had been counted. This was repeated for a total of 
9 sporangia. 



RESULTS 
Branch fecundity and mortality 

The frequency of branching, or dichotomy, declines with increasing segment position, which 
is equivalent to increasing age of the whole plant, in all three populations. This results 
in the opposite of a tree-like structure (Fig. 3). There is a much lower frequency of branching 
in the Svalbard population in the first year. The Snowdonia population has a significantly 
higher dichotomy frequency than the Abisko and Svalbard populations in the fourth and 
fifth years. Overall the population which has much the lowest dichotomy frequency of 
the three populations is that from Abisko. 

The mortality of branches is generally below 10% in the first five years of growth (Fig. 4). 
After this period mortality is generally highest in the Abisko population. 



MODULAR GROWTH OF HUPERZIA SELAGO 



365 



c 
a) 

c_> 

c_ 

O 
DL 



50- 
45- 
40- 
35' 
30- 
25 
20 
15 
10 
5 





4 5 6 7 
Segment position 



10 



FIGURE 4. Changes in mean branch mortality (%) with increasing distance from the base of Huperzia 
selago plants from Snowdonia (circles), Abisko (triangles) and Svalbard (squares). 





110-] 




100- 




90- 


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20 
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4 5 6 7 8 9 
Segment position 



10 11 12 13 



FIGURE 5. Changes in mature segment weight with increasing distance from the base of Huperzia 
selago plants from Snowdonia (circles), Abisko (triangles) and Svalbard (squares). Means with S.E. 
bars shown. 



366 



FERN GAZETTE VOLUME 13 PART 7 (1990) 



Growth of branches 

The weight of segments is affected by a) the population they belong to, b) their position, 
c) their age and d) if they have dichotomised. Segments of material from Abisko weigh 
less than those from the other two sites (Fig. 5), while the segments of the Snowdonia 
population are by far the heaviest. In general, the basal segments are heavier than those 
higher up the plant (Fig. 5). As they age, dry weight is accumulated over one year in 
material from Snowdonia, whereas segments increase in dry weight over a 1-8 year period 
in material from Abisko and Svalbard (Fig. 6). 

Larger segments are associated with dichtomies and segments with a dichotomy in material 
from Snowdonia were 72% heavier (55 mg segment ~ 1 ) than those without (32 mg 
segment ~ 1 ). There is however, an interaction between segment position, and age, and 
dichotomy which has not been quantified here. 



45- 



40- 



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35 


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12 3 4 5 6 7 8 9 

Segment age (yeans) 

FIGURE 6. Changes in unbranched segment weight with increasing age in Huperzia selago plants 
from Snowdonia (circles), Abisko (triangles) and Svalbard (squares). Data is for segments of all positions. 
Means with S.E. bars shown. 



Allocation of dry weight to roots 

The proportion of total plant weight allocated to roots declines significantly from 
24.6% ± 1.7% in one year old plants to 9.8% ± 0.8% in two or more year old plants 
from Snowdonia (t=8.879, d.f.= 18, p< 0.001). There is no significant effect of plant age 
on the percentage biomass in the roots of 2 or more year old plants (r=0.058, d.f.= ll, 
p >0.05). 



Total plant weight 

There appears to be a biphasic growth pattern in total dry weight of plants for the Snowdonia 
and Svalbard populations (Fig. 6). The initially more rapid increase in dry weight (RGR 



MODULAR GROWTH OF HUPERZIA SELAGO 



367 



= 2.23 and 1.76 g g" 1 y 1 , calculated from data in Fig. 7, for the Snowdonia and Svalbard 
populations, respectively) is in the first two years of growth. The slower RGR after this 
is very low (0.375 and 0.136 g gr 1 y~ 1 ). The total dry weight of the H. selago plants from 
Abisko showed no significant correlation with age. 

Vegetative reproduction 

As branches of H. selago grow apically, the basal segements become decumbent and as 
a result the plant spreads out either centrifugally on flat terrain or downhill on a slope. 
Fragmentation of a large plant in theory is possible, but large clones of H. selago are 
rare, with clones 1-2 metres across having only been reported from the undisturbed coniferous 
forests of Russia (Syelivanova-Gorodkova 1968). 

The normal mode of reproduction is by bulbils, as all plants harvested or examined 
possessed the distinctively shaped leaves of the bulbil at the very base of the plant. The 



2- 



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= -1H 
O 

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4 


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8 


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12 


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14 


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16 



PLANT AGE (years) 



FIGURE 7. Relationship between whole plant weight (expressed on a logarithmic scale) and age of 
Huperz/'a selago plants from Snowdonia (solid circles and line) and Svalbard (open circles and dashed 
line). Means with S.E. bars shown for three or more replicates. 



368 



FERN GAZETTE VOLUME 13 PART 7 (1990) 



production of bulbils does not normally start until the third or fourth year of the plant's 
life (Fig. 8) and they are therefore, not released until the fourth or fifth year. The number 
of bulbils produced per segment increases with the age of the plant (Fig. 8). Despite the 
branch segments of the Abisko populations being lighter and hence smaller, the number 
of bulbils produced per segment is greater than that produced by the branch segements 
of the Snowdonia population over the first 8 years (Fig.7). There is however, a much higher 
bulbil production per segment in the Snowdonia population later in the life of the plant 
(Fig.8). 

The mean weight of bulbis produced by the H. selago plants from Svalbard is intermediate 
(1.04 ± 0.06 mg) between those of the Abisko population (0.69 ± 0.04 mg) and the Snowdonia 
population (1.88 ± 0.10 mg). 



en 

CD 
CD 

c_ 
c 



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o 

c 

CL 



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=3 
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10 
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6 
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4- 
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Segment position 



FIGURE 8. Changes in bulbil production with increasing distance from the base of Huperzia selago 
plants from Snowdonia (circles) and Abisko (triangles). Means with S.E. bars shown. 



Spore production 

Larger segments, which naturally weigh more, produce more sporangia (Fig. 9). The allocation 
to spores is 7.0% ± 1.2% of the dry weight of the one-year-old segment which produces 
them. The number of spores in each sporangium is 19,000 ± 2,000. This means that a 
plant from Snowdonia, will produce on average 0.97 x 10 6 spores segment" 1 . However the 
viability of spores is only 6.7% ± 2.2%. 



DISCUSSION 
H. selago is very widespread and frequently a component of montane, boreal and tundra 
environments, although it is never dominant in the communities in which it occurs (Headley 
1986). It is a plant of very low relative growth rate and is consequently found growing 
in infertile habitats or niches within these environments (Headley 1986). 



MODULAR GROWTH OF HUPERZIA SELAGO 369 



There is a very strong pattern of basal (positional) branching in H. selago, which does 
not differ very greatly between the populations sampled in this study or elsewhere (Turmel 
1982). As H. selago does not have secondary thickening the basal branching which is the 
opposite to a tree-like structure, maximises the support of higher branches. However, the 
support is inefficient and individual branches gradually become decumbent at the base as 
the weight of a growing branch exceeds the capacity of the lower part of the stem to 
support it (Niklas & O'Rourke 1982). This pattern of basal branching is also observed 
in the vertical branch systems of the stoloniferous members of the Lycopodiaceae; L. 
annotinum and L. clavatum (Callaghan et al 1986a). 

The annual weight increments of individual branches are generally very low, and are 
very similar to those of other evergreen tundra plants (4 to 62 mg y~ 1 ) at Eagle Summit, 
Alaska (Miller 1982). The climatic conditions experienced on the top of Mount Njulla, 
are more severe than those in Snowdonia and mean segment weight is on average 77% 
± 3% lower in plants from Njulla compared to that of Snowdonia plants. Although climatic 
conditions affect segment weight, and age also affects the weight of developing segments, 
the position of the segment on the plant is the most important determinant of segment 
weight within any one plant. 

H. selago differs from L. annotinum in not having a foraging strategy, but a tolerance 
of very stressed climatic and edaphic environments. H. selago is found as far north as 
Ellesmere Island and northern Greenland (82°N) and is found in soils that are naturally 
impoverished of major plant nutrients (0.4-18.2 ppm N; 4.4-5.8 ppm P). This is one reason 
for the very low RGR. The plant can respond to low phosphorus supply by increasing 
the rates of phosphate uptake (Headley 1986). However the plant can maintain much of 
its annual nutrient requirements by efficient internal recycling of the elements from basal 
segments to apical segments (50-80% for nitrogen and 40-60% for phosphorus) (Headley 
1986). The immobility of H. selago compared to that of L. annotinum means that the 
plant has to be more efficient at capturing and utilising nutrients as root growth is restricted 
spatially. As a consequence, there is a much larger allocation of dry weight to roots in 
H. selago (10% of total plant weight) than in L. annotinum (5% of total plant weight). 

Very old plants can become well established and have potentially indefinite growth in 
sheltered undisturbed forest habitats in Russia (Syelivanova-Gorodkova 1968). H. selago 
is however normally short lived (6-16 years) in the open habitats in Europe due to the 
premature death of the branches. As this species, like all the temperate members of the 
Lycopodiaceae, very rarely produces adventitious shoots, the removal of the branch tips 
by grazing or burning, as often happens in upland Britain, kills the plant. The plant may 
also show signs of premature death due to an inability to produce new roots as a consequence 
of being rooted in small crevices in rocks. 

H. selago overcomes the problems of a limited life-span and immobility through successful 
reproduction by bulbils. H. selago plants may produce large quantities of bulbils over their 
whole life span, for example a plant from Snowdonia produced a total of 1166 bulbils 
over its 13 year growing period, which represented an investment of 18% of its dry weight 
to this form of reproduction. The allocation of this amount of dry weight to vegetative 
reproduction by bulbils is very similar to that observed for Saxifraga cernua and Polygonum 
viviparum (Wehrmeister & Bonde 1977; Petersen 198 1 ). Reproduction by bulbils is particularly 
successful in the sparse vegetation of severely stressed and disturbed environments in the 
arctic (Bell & Bliss 1980; Callaghan & Emanuelsson 1985). 

There is normally 100% germination of fresh bulbils which readily established on a 
wide range of substrata, including raw humus, peat, moss cushions, cracks in rocks, sand 
and rankers. This is typical for other species that reproduce by bulbils and plantlets (Harmer 
& Lee 1978b). This mode of reproduction is adequate for maintaining a viable population 
and spreading it within a particular site, especially a moderately closed community. The 



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FERN GAZETTE VOLUME 13 PART 7 (1990) 



large nutrient reserves within the bulbils (Smith 1920; Harmer & Lee 1978a; Headley 1986) 
are sufficient to enable the plant to grow up through competing grasses in the uplands 
of North Wales, the Lake District and Scotland, while opportunistic germination maximises 
the use of short growing seasons. 

The distance between safe microsites are too great for L. annotinum to cross by means 
of subsidised growth of the stolon system in the extremely patchy environments of fell- 
fields, screes and mountain tops, and rhizomatous/stoloniferous plants cannot generally 
tolerate the disturbance associated with some of these habitats. It is here however, that 
H. selago and other bulbiferous/viviparous species have an advantage. The bulbils can 
successfully be transported by wind and water. This is a more opportunistic method of 
vegetation reproduction. 



100 




10 20 30 40 50 60 

SEGMENT DRY WEIGHT (mg) 



FIGURE 9. Relationship between the number of sporangia per segment and segment dry weight in 
Huperzia selago plants from Snowdonia. r = 0.718, P 0.001. 



The low allocation of dry weight to sexual reproduction ( 7.0%) and the absence of 
genets in closed vegetation is similar to that for L. annotinum and other perennials of 
the tundra (Chester & Shaver 1982; Callaghan & Emmanuelsson 1985). H. selago can only 
successfully spread any significant distance by the numerous spores produced in the la